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Bulletin of the 

British Museum (Natural History) 



Spectacles and Silver Ys: a synthesis of the 
systematics, cladistics and biology of the 
Plusiinae (Lepidoptera: Noctuidae) 

Ian J. Kitching 



Entomology series 

Vol 54 No 2 28 May 1987 



The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four 
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London SW7 5BD, 
England. 



World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) 



i Trustees of the British Museum (Natural History), 1987 



The Entomology series is produced under the general editorship of the 

Keeper of Entomology: Laurence A. Mound 

Assistant Editor: W. Gerald Tremewan 



ISBN 565 06025 2 

ISSN 0524-6431 Entomology series 

Vol 54 No 2 pp 75-261 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD Issued 28 May 1987 



Spectacles and Silver Ys: a synthesis of the 
systematics, cladistics and biology of the Plusiinae 
(Lepidoptera: Noctuidae) 

Ian J. Kitching 

Department of Entomology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Contents 

Synopsis 75 

Acknowledgements 76 

Introduction 76 

The history of the subfamily Plusiinae 77 

The higher classification of the plusiines 77 

The position of the Plusiinae within the Noctuidae 92 

Taxa examined 93 

Terminology 94 

Methods 97 

Characters 97 

Analytical procedure 112 

The 648-step cladograms 113 

1 . Agrapha and Plusiopalpa 113 

2. Plusidia 115 

3. Autographa, Cornutiplusia and Plusia 115 

4. Loboplusia 117 

5. Rachiplusia and the branching sequence of the tribes and subtribes 117 

6. Outgroups 119 

The 647-step cladograms 121 

Evaluation of the preferred cladogram 121 

Character analysis 124 

Clade analysis 138 

Intra-outgroup analysis 167 

Consensus classification 170 

Analytical procedure 170 

Results 170 

The positions of Shensiplusia, Desertoplusia and Pseudochalcia 171 

The higher classification of the Plusiinae 174 

Comparison with previous classifications and phylogenies 175 

Discussion 177 

Larval foodplant associations 177 

Biogeography 178 

Conclusions 179 

Addenda: notes added in proof 180 

References 180 

Appendix 1. Genera and type species of Plusiinae and outgroups examined 187 

Appendix 2. Plusiinae generic-level taxa not employed in this study 190 

Appendix 3. Data matrix 191 

Appendix 4. New taxa and synonymy, status changes and revivals 195 

Appendix 5 . Generic classification of the Plusiinae 195 

Index 259 

Synopsis 

A comprehensive historical review of the higher classification of the Plusiinae is presented, with particular 
reference to the different generic concepts and interrelationships that have been proposed. The position of 

Bull. Br. Mus. nat. Hist. (Ent.) 54 (2): 75-261 Issued 28 May 1987 



76 IAN J. KITCHING 

the subfamily within the Noctuidae is considered separately. A cladistic analysis of 47 generic-level taxa of 
plusiines and 11 outgroups was performed, using the type species of each as an exemplar. A total of 216 
characters is described, drawn from the morphology of the head, prothorax, vestiture, wings, legs, 
abdomen and male and female genitalia. These data were analysed using the numerical cladistic program 
PAUP and at least 50 equally-parsimonious cladograms of 648 steps were generated. The character state 
changes supporting the various alternative subtopologies are discussed and a preferred 648-step cladogram 
constructed. A further analysis revealed the existence of 16 647-step cladograms but, after character 
analysis, these were all rejected. The preferred 648-step cladogram is evaluated, first in terms of 
characters, and then in terms of its constituent clades. Under each of the genera in the latter section , a brief 
review of biology, distribution and other points of interest is also included. This assessment allowed 
characters of low reliability to be identified and subsequently eliminated from the data. The reduced data 
set, less the outgroups, was analysed once more using PAUP. At least 50 equally-parsimonious 
cladograms were again found. A manual strict consensus cladogram analysis was then performed using 
these cladograms and, from the results, a consensus classification of the Plusiinae constructed, using the 
conventions for annotated Linnaean hierarchies. Four tribes are recognized in the Plusiinae: the Omorphi- 
nini, Abrostolini, Argyrogrammatini and Plusiini. The last is divided into three subtribes: the Euchalciina, 
Autoplusiina and Plusiina. Several genera (Shensiplusia, Pseudochalcia, Diachrysia, Anagrapha, Rachi- 
plusia, Loboplusia and Erythroplasia) cannot be placed with confidence and are included incertae sedis in 
the classification. Cornutiplusia, Lophoplusia, Ctenoplusia and Thysanoplusia are recognized as good 
genera, although with reservations regarding the last of these. Pseudoplusia, Eutheiaplusia, Acanthoplu- 
sia, Adeva and Caloplusia are interpreted as subgenera of Chrysodeixis, Plusiotricha, Ctenoplusia, 
Euchalcia and Syngrapha respectively. This classification is compared with the five recent higher 
classifications proposed for the Plusiinae. Finally, the larval foodplant associations and biogeography of 
the subfamily are briefly discussed. One new subtribe is erected and five new combinations made. One 
species and one generic synonymy are proposed. Four previously synonymized genera are recognized as 
valid while a further five genera are reduced to subgeneric status. 

Acknowledgements 

This study was carried out during the tenure of a Junior Research Fellowship awarded by the Trustees of 
the British Museum (Natural History), which is gratefully acknowledged. I thank my colleagues at the 
British Museum (Natural History) for their support and comments, the staff of the Electron Microscope 
Unit, BMNH, for their assistance, and the staff of the Photographic Unit, BMNH, for the photographs 
reproduced in this paper. I also thank Dr R. de Jong, Rijksmuseum van Natuurlijke Historic Leiden, and 
Dr W. Dierl, Zoologische Staatssammlung, Munich, for the loan of type specimens. 

Introduction 

For over 80 years, the higher classification of the family Noctuidae has consisted of a 
non-hierarchical series of largely artificial subfamilies, based upon a small number of simple and 
readily observable characters. There have been frequent small adjustments to the positions of 
individual genera; more rarely a new tribe has been erected (although usually defined only for a 
regional fauna), and never has a whole subfamily been revised. However, if a useful and 
predictive higher classification of the noctuids is to be constructed, then problems have to be 
addressed at the tribal level and above. 

Subsequent to my recent review of noctuid higher classification (Kitching, 1984), I decided to 
analyse the interrelationships of the genera of the subfamily Plusiinae, by the critical application 
of cladistic methodology . The plusiines were chosen for a number of reasons. First , and perhaps 
foremost, there were reasonable grounds for supposing that, even as currently constituted, the 
Plusiinae are a monophyletic group. Thus, it would be unlikely that any of the constituent genera 
would be overlooked by their being placed in other subfamilies. This supposition is strongly 
supported by the observation that, since Hampson's (1913) Catalogue, no pre-existing genera 
have been transferred into the Plusiinae from elsewhere in the Noctuidae. Additional genera 
have arisen either by the discovery of new species (e.g. Loboplusia, Eutheiaplusia) or by the 
subdivision of pre-existing plusiine genera. A few genera (e.g. Phyprosopus Grote, Diloba) 
have been transferred out of the Plusiinae. Second, the plusiines are a relatively small subfamily 
of around 400 species and 45 genera. They were thus more amenable to analysis within my terms 
of reference than, say, the more economically important but much larger Noctuinae (with 



SYSTEMATICS OF THE PLUSIINAE 77 

approximately 2000 species in 150 genera). This is not to say that the plusiines are merely of 
academic interest. The subfamily includes a number of major agricultural pests, notably 
Trichoplusia ni, Pseudoplusia includens, Chrysodeixis spp. and Thysanoplusia orichalcea 
(Fabricius). Thus, a predictive higher classification of the Plusiinae is vital for the effective 
applied entomology of the group. Finally, the collections of the British Museum (Natural 
History) (BMNH) contain specimens of about 80% of the world species and 96% of the genera 
of plusiines: relatively complete coverage by any standards. 

It was impossible to conduct a species-level analysis of the subfamily and so some selection of 
taxa to be included had to be performed initially. I decided to concentrate on one species from 
each genus or genus-group taxon, choosing the type species for convenience. Characters were 
drawn primarily from the external morphology of the adult moths. Little consideration was 
taken of the internal morphology, and none of the immature stages or biology. A cladistic 
method was chosen primarily because of the explicit manner in which it handles characters and 
their state transformations. There is no excuse for hiding behind vague or authoritarian 
statements of relationships. I hope my reasoning on such issues as character coding, polarity and 
interpretation are clear: that of many previous authors has not been so. 

The history of the subfamily Plusiinae 

The higher classification of the plusiines 

In the following historical review of the Plusiinae, several terminological conventions have been 
employed. Members of the subfamily in general are referred to as 'plusiines', while 'plusias' 
refers collectively to the tribes Argyrogrammatini and Plusiini (i.e. the Plusiinae less the 
Abrostolini and Omorphinini). The taxon names used are those employed in the publication 
being discussed at that point. Current nomenclature, where this differs, is given in square 
brackets. This approach was adopted to facilitate back-referencing by the reader and to clarify 
some of the earlier nomenclatural anarchy. 

The emergence of the Plusiinae 

Linnaeus (1758) described the first plusiine noctuids within the subsection of Phalaena Noctua 
characterized as 'spirilingues dorso cristato'. Of these, triplasia was well separated from the 
remaining five species (chrysitis, gamma, interrogationis, jota Linnaeus and festucae). This 
arrangement was followed by Denis & Schiffermuller (1775), who included triplasia (together 
with asclepiadis [Denis & Schiffermuller] and two other noctuids) in the informal group Y 
(Larvae Pseudogeometrae, Phfalaenae] Noctuae Submetallicae), and the other plusiines in 
group Z (Larvae Semigeometrae, Phfalaenae] Noctuae Metallicae). These groups were 
accepted by Borkhausen (1792) and formally named by Ochsenheimer (1816) as the genera 
Abrostola and Plusia. Haworth (1809), however, chose to treat triplasia and asclepiadis as a 
section of Noctua (19 Arctuo-strigatae) and included the other plusiines in the genus Phytometra 
(section Metallicae), a decision that was to have important repercussions subsequently. 

Hiibner [1821] increased the number of plusiine genera to ten. Abrostola was again placed in a 
separate 'family', while the remaining plusiine species were allocated to Euchalcia, Agrapha, 
Syngrapha, Autographa, Polychrysia, Panchrysia, Diachrysia, Chrysaspidia and Chrysodeixis. 
However, these genera were , by and large , ignored for the remainder of the nineteenth century. 

The Plusiinae as a family-group entity originated as the Tribus Plusidi, including three genera: 
Abrostola, Chrysoptera [Lamprotes] and Plusia (Boisduval, 1828). Duponchel (1844) modified 
this arrangement by treating the latter two groups as 'subgenera' of Plusia. Plusia (s. str.) was 
further subdivided into three unnamed sections based on wing pattern. Guenee (1841; 1852-4) 
and Herrich-Schaffer (1845) proceeded along similar lines. 

This bigeneric system persisted for many years (e.g. Grote, 1874; 1875; Hampson, 1893-5), 
although Smith (1882-3) united the two into a single genus Plusia. Gradually, however, opinion 
swung towards the view that the omnibus genus Plusia ought to be subdivided, given the 
structural diversity displayed by its included species. Both Grote (1882) and Tutt (1891-2) 



78 IAN J. KITCHING 

considered Deva [Pseudeva] to be a good genus, and by 1875, Grote had accepted Hiibner's 
[1821] generic groupings, to which he added Chrysanympha. Butler (1892) also argued for 
separate generic status for Polychrysia and considered Deva (sensu Grote and others) to be 
synonymous. Polychrysia thus included additional species currently referable to Lamprotes and 
Pseudeva. 

On the basis of characters drawn from the wing-shape, labial palps and eyes, Dyar (1902) 
recognized seven genera of plusias: Polychrysia, Panchrysia, Plusia, Autographa, Euchalcia, 
Syngrapha and a new genus, Eosphoropteryx. This arrangement was endorsed, for the most 
part, by Ottolengui (1902) in a paper published in the same journal only a few pages before that 
of Dyar. Forbes (1914) reverted to the omnibus Plusia but included Dyar's subdivisions in a 
note. 

Meanwhile, Hampson (1913) was subdividing the plusiines along completely different lines. 
The subfamily was renamed the Phytometrinae, with the type genus and species, Phytometra 
festucae. This change resulted from Hampson's criterion for type species designation. Irrespec- 
tive of previous designations, Hampson chose as the type species of a genus, the first species 
included when that genus was described, unless otherwise explicitly specified by the author at 
the time (McDunnough, 1916). Thus, the type species of Phytometra Haworth was considered to 
be festucae, despite Westwood's (1840) designation of aenea ([Denis & Schiffermuller]) [= 
viridaria (Clerck)]. Thus Plusia, the type species of which was taken by Hampson to be deaurata, 
became a subjective synonym of Phytometra. 

In addition to 'Phytometra' , fourteen other genera were placed in the 'Phytometrinae', on the 
basis that they were quadrifine noctuids with lashed eyes. In his 'phylogeny' of the subfamily, 
Hampson considered that the unknown ancestor had given rise to Omorphina (transferred to 
the Erastriinae by Kostrowicki, 1961; but see below), Diloba (of current uncertain position, but 
probably a noctuid), Abrostola and Palaeoplusia. Mouralia was derived from Abrostola, while 
Palaeoplusia was ancestral to 'Phytometra . This last genus contained by far the majority of the 
plusiines but from it arose five further genera: Plusidia, Pseudeva, Eosphoropteryx, Chrysoptera 
[Lamprotes, Polychrysia] and Syngrapha. Plusiopalpa was considered to be a derivative form of 
Chrysoptera, while, by additional tibial spination on the fore- and mid-legs and a reduction in the 
size of the eyes, Autographa (comprising only the present Syngrapha parilis (Hiibner), another 
result of Hampson's type species designations), Rachiplusia and Caloplusia were derived from 
Syngrapha. Syngrapha itself was divided into two parts, corresponding approximately to the 
current genera Syngrapha and Autoplusia. In Chrysoptera, sica (Graeser) was distinguished 
from the remaining species by an excised forewing termen, while 'Phytometra' was split into four 
groups, depending on whether the male antennae were bipectinate and apically ciliate ('P.' 
pterylota), bipectinate and apically serrate ('P.' psectrocera Hampson), serrate ('P.' lavendula 
(Hampson) and 'P.' gerda (Piingerler)) or ciliate (the remaining 151 species). Chrysanympha 
formosa was placed in Abrostola. 

Hampson's arrangement was followed in its entirety by the authors of 'Seitz': Warren (1913), 
Draudt (1940) and Gaede (1937-9). However, other workers were far from satisfied. McDun- 
nough (1916) corrected many of Hampson's erroneous type species designations, including that 
of Phytometra. This allowed him to replace 'Phytometra' with Autographa, return parilis to 
Syngrapha and to reinstate Plusiinae as the subfamily name. At this time, it should be noted, the 
type species of Plusia was considered to be chrysitis (Linnaeus), a point to which I shall return 
below. 

McDunnough' s revision of the N ear ctic plusias 

Barnes & McDunnough (1917) advocated a system that was something of a hybrid between 
those of Dyar (1902) and Hampson (1913), and which was largely accepted by McDunnough 
(1938) in his later checklist. However, shortly afterwards, McDunnough (1944) published a 
revision of the Nearctic plusias, which represented a significant departure from all previous 
classifications, and for the first time in plusiine systematics, genitalic characters were employed. 
The subfamily (less Abrostola and Mouralia) was split into three sections based on the form of 



SYSTEMATICS OF THE PLUSIINAE 79 

the clavus. Section A, in which the clavus was a short, slightly setose knob arising from a broad 
base, included Syngrapha, Anagrapha and Caloplusia, and was additionally characterized by 
'readily recognizable' tibial spines. Section C comprised Pseudeva and Adeva, genera in which 
the clavus was not clearly defined, while section B, in which the clavus was a long, thin, variably 
setose rod, contained the balance of the species. This latter section comprised twelve genera: 
Autoplusia, Trichoplusia, Argyrogramma, Pseudoplusia, Rachiplusia, Chrysaspidia [Plusia], 
Autographa, Chrysanympha, Eosphoropteryx, Polychrysia, Plusia [Diachrysia] and Agrapha 
[Allagrapha]. Palaeoplusia was synonymized with Chrysaspidia [Plusia]. 

By way of an aside, it was subsequently noted by Franclemont (1964) that Grote (1896) had 
designated Phalfaena] Noct[ua] glauca (Cramer) [= ahenea Hiibner] as the type species of 
Agrapha. Agrapha originally contained two species, aerea and glauca, which Franclemont 
considered were not congeneric. Because he agreed with McDunnough that aerea represented a 
genus distinct from Plusia [Diachrysia] but that Agrapha was inappropriate, Franclemont 
proposed the genus Allagrapha, noting similarities in the male and female genitalia with 
Macdunnoughia that were not shared with Plusia [Diachrysia]. 

Forbes (1954) followed McDunnough's (1944) subdivision of the plusiines but treated the 
latter's genera as subgenera of Plusia , as did Zimmerman (1958) . Inoue & Sugi ( 1958) attempted 
to apply the new system to the Japanese fauna but the result was not very satisfactory; 
Autographa, in particular, became highly heterogeneous. 

Meanwhile, the studies of Forbes (1910), Fracker (1915), Gerasimov (1935) and Hinton 
(1946), amongst others, were stimulating interest in the use of the detailed larval morphology in 
lepidopteran classification. Initially, largely due to lack of material, the results consisted mainly 
of descriptions (e.g. Gardner, 1941; 1947; 1948a) but soon sufficient taxa had been examined 
and described to allow some comparative work to be undertaken. Crumb (1934) split the 
plusiine larvae into two groups based on the ordination of the crotchets. Gardner ( 1948a) noted 
differences in chaetotaxy, skin ornamentation, proleg number, maxillary palp and antennal 
segment shape but made no subdivisions using these characters. A similar dichotomy in 
chaetotaxy was found by Timlin (1955) but unfortunately, like Gardner, the number of species 
examined was too small to be able to conclude anything definitive about the interrelationships of 
the genera. 

This position was changed markedly by Crumb's (1956) study of the larvae of 25 species in 14 
genera of Plusiinae. On the basis of characters drawn mainly from the mandibles and chaeto- 
taxy, he grouped the plusiines into nine informal genera. Four of these (Abrostola, Polychrysia, 
Chrysaspidia [Plusia] and Caloplusia) corresponded to McDunnough's generic concepts. 
Trichoplusia, Pseudoplusia and Argyrogramma were placed in a single taxon, as were Autoplu- 
sia and Rachiplusia, and Plusia [Diachrysia] and Agrapha [Allagrapha]. Those species of 
Syngrapha (rectangula (Kirby) and celsa (Edwards)) that possessed SV2 on segments Til and 
Till, and which also fed as larvae on conifers, were separated as 'Genus no. 11', while those 
Autographa that possessed SV2 on segment Al (mappa (Grote & Robinson), ampla (Walker) 
and corusca (Strecker)) were included in Syngrapha. Autographa was thus restricted to those 
species lacking SV2 on segment Al - biloba (Stephens), californica (Speyer), gamma and 
precationis (Guenee). 

Beck (1960) examined the larvae of several European genera of Plusiinae: Abrostola, Plusia 
[Diachrysia] , Autographa, Chrysaspidia [Plusia], Syngrapha [Euchalcia] and Polychrysia, but 
also drew no conclusions as to possible affinities. 

Although these early larval studies made relatively few hard statements regarding the higher 
classification of the Plusiinae, they did demonstrate that larvae could be of use taxonomically 
and paved the way for future research. 

Kostrowicki and the Palaearctic plusias 

The next major revision of the Plusiinae was performed by Kostrowicki (1961), who studied the 
Palaearctic fauna, comprising 83 species in 23 genera. Abrostola and Mouralia were omitted 
from consideration . Kostrowicki was generally in favour of McDunnough's ( 1944) arrangement 



80 



IAN J. KITCHING 



of the Nearctic fauna but expressed reservations regarding its uncritical application to the 
Holarctic fauna. Forbes' (1954) reduction of McDunnough's genera to subgeneric status was 
rejected because this would 'introduce quite unnecessary inequalities with the other subfamilies 
of [Noctuidae]'. Crumb's (1956) arrangement found more favour but Kostrowicki questioned 
both the synonymy of Trichoplusia, Pseudoplusia and Argyrogramma, and the reality of 
Crumb's 'genus no. 11'. 

Kostrowicki then proposed his own phylogeny of the plusias, which he hoped would correct 
some of McDunnough's and Crumb's parochial errors (Fig. 1). Argyrogramma was considered 
to be the ancestral plusia lineage, which gave rise to three groups of genera. The first consisted of 
Anadevidia, Pseudoplusia, Trichoplusia, Chrysodeixis, Macdunnoughia and Diachrysia [Thy- 
sanoplusia] . This last genus, for some reason, was omitted from the phylogenetic tree. The 
second group was characterized by a tendency for the palps to be rather longer than in the other 
plusiines. One subgroup, Plusia [Diachrysia] , Polychrysia and Panchrysia, was considered to be 
possibly related to Diachrysia [Thysanoplusia] '. The second subgroup consisted of a postulated 




Caloplusia 



Fig. 1 Phylogenetic tree of Kostrowicki. Originally, his tree depicted supposedly ancestral genera 
(Argyrogramma, Diachrysia (as Plusia), Panchrysia, Euchalcia, Plusia (as Chrysaspidia) and Auto- 
grapha) by thicker branches and branch lengths appear to have been related in some way to degree of 
anagenetic change. These two aspects were eliminated and only the topology retained in the above 
figure. Generic names have been changed into their modern forms. (Redrawn from Kostrowicki, 1961.) 



SYSTEMATICS OF THE PLUSIINAE 81 

ancestral genus, Euchalcia, and three derivative genera, Plusidia, Chrysoptera [Lamprotes] and 
Pseudeva. Kostrowicki commented on Eosphoropteryx in connection with these genera but felt 
he could not place it within the group because of the distinctive appearance of the genitalia. 

The third group to be derived from Argyrogramma was based around Autographa and 
Chrysaspidia [Plusia] . The latter was considered to have given rise to Chrysanympha, while the 
former had given rise to two groups of genera, both characterized by spinose tibiae. The first 
consisted of the Nearctic Autoplusia and Rachiplusia and the Old World genus Cornutiplusia . 
The second included the primarily arctic-alpine genera Syngrapha and Caloplusia, and the more 
widely spread Nearctic Anagrapha, of which Syngrapha was considered to be the ancestral 
group. Kostrowicki rejected Crumb's (1956) delimitations of Autographa and Syngrapha 
because the divisions based upon the larvae were not reflected by differences in the genitalia. 

Kostrowicki's scheme showed several important discrepancies compared to the generic 
interrelationships that had been proposed previously. Several were due to incorrect assignment 
of species to genera, which then became structurally heterogeneous, or to straightforward 
misidentification (Ichinose, 1963a; Dufay, 1970a). Into the first category fell, for example, the 
placing of Autographa aemula ([Denis & Schiffermuller]), A. bractea and A. argyrosigna 
(Moore) in Chrysaspidia [PlusiaJ. The inclusion of Antoculeora ornatissima and Erythroplusia 
rutulifrons in Argyrogramma, and Erythroplusia pyropia (Butler) in Anadevidia were due to 
misidentifications. The male genitalia of Antoculeora ornatissima figured have the quadrate 
saccus typical of Argyrogramma and are probably those of Argyrogramma signata (Fabricius) 
(Dufay, 1970a: 150, footnote), the only species of Argyrogramma (s. str.) to occur in eastern 
Asia. 'Argyrogramma rutulifrons' appears to be very closely related to Argyrogramma circums- 
cripta (Freyer), and may even be that species. The identity of 'Anadevidia pyropia' remains a 
mystery. 

The rising influence of larvae: Ichinose's revisions 

Following on from a series of studies of plusiine larval morphology (Ichinose, 1958a; b; 1960a; 
1962a), Ichinose (19626) produced a revision of the Japanese Plusiinae, contemporaneous with 
Kostrowicki's work. Ichinose considered that McDunnough had oversplit the subfamily and that 
many of the characters employed were of little taxonomic significance, particularly the form of 
the clavus. Instead, Ichinose preferred the generic assignments suggested by Crumb (1956), 
although he felt that Crumb had placed too much emphasis on certain characters and thus had 
reached some dubious conclusions. Although Ichinose was in favour of using characters drawn 
from all stages of the life-history in the construction of classifications, at the generic level he 
predominantly used larval characters, particularly from the chaetotaxy. 

Ichinose recognized twelve genera of Plusiinae. The delimitation of eight of these was 
uncontroversial and conventional: Abrostola, Plusidia, Chrysoptera [Lamprotes], Euchalcia, 
Podioplusia [Anadevidia], Chrysaspidia [Plusia] and Syngrapha. Autographa was divided into 
three subgenera: Autographa (s.str.), Scleroplusia [Macdunnoughia (s.l.)] and Erythroplusia, 
separated on the basis of the divergent form of the male genitalia but united by a uniform larval 
morphology. Ichinose, like Kostrowicki, separated intermixta from the remaining species of 
Plusia [Diachrysia] and placed it, together with daubei (Boisduval) and ochreata, in Diachrysia 
[Thysanoplusia] . The golden wing pattern of intermixta and the genus Plusia [Diachrysia] was 
considered to be convergent. 

While this reallocation of species left Plusia [Diachrysia] more compact and homogeneous, 
the three species of Diachrysia [Thysanoplusia] showed diverse possible affinities. Overall, 
Ichinose considered Diachrysia [Thysanoplusia] to be much more closely related to Chry- 
sodeixis than to Plusia [Diachrysia] . However, while the chaetotaxy and skin ornamentation of 
intermixta suggested further affinities with Podioplusia [Anadevidia] and Euchalcia, the chaeto- 
taxy and external appearance of daubei suggested a possible connection with Chrysodeixis. 

With regard to Chrysodeixis itself, Ichinose followed Crumb (1956), including Trichoplusia 
(and also the species now assigned to the nominal genera Ctenoplusia (albostriata) and 
Acanthoplusia (agnata (Staudinger) and tarassota)). 



82 IAN J. KITCHING 

Ichinose presented his overall view of the generic relationships in the form of a 'phylogenetic 
tree' (Fig. 2). Exactly how this dendrogram is to be interpreted is uncertain. It agrees reasonably 
well with Kostrowicki's tree (Fig. 1), in that the Polychrysia-group and the Autographa-group 
arose independently from a Chrysodeixis-Argyrogramma-type ancestral form. In both treat- 
ments, Podioplusia [Anadevidia] was of uncertain position, while Ichinose's clarification of the 
positions of Macdunnoughia and Erythroplusia represented a considerable improvement over 
Kostrowicki's placings. 

Meanwhile, Claude Dufay had been busily publishing a large number of studies on plusiine. 
alpha-taxonomy. He eventually ventured his opinions on the higher classification of the 
subfamily in his revision of the Madagascan fauna (Dufay, 1970a). Following the obligatory 
discussion of Hampsonian type species designations, Dufay proceeded with a brief discussion of 
the studies of McDunnough, Kostrowicki and Ichinose. Like Ichinose (1963), Dufay disagreed 
with some of Kostrowicki's generic assignments, particularly regarding Argyrogramma. How- 
ever, he also considered Ichinose's treatment to be unsatisfactory, for although the groupings 
were certainly more natural than those of Kostrowicki, several genera (in particular Chryso- 
deixis) were too heterogeneous. Dufay preferred to keep Trichoplusia, Argyrogramma and his 
own Ctenoplusia separate from Chrysodeixis. 

Like Kostrowicki, Dufay's revision was also reviewed in Japan (Sugi, 1972). The major 
criticism appeared to be Dufay's inclusion of [Thysanoplusia] in Trichoplusia, which thereby 
rendered that genus heterogeneous in terms of larval chaetotaxy and skin ornamentation. 

Shortly afterwards, Ichinose (1973) published an upgraded classification of the Japanese 
plusiines. Autographa now included Macdunnoughia and Sclerogenia as subgenera but the 
previously included subgenus Erythroplusia was elevated to full generic status. Erythroplusia 
comprised two subgenera, Erythroplusia (s.str.) and Antoculeora, united by characters of the 

P Abrostola 



/ * 


Chrysodeixis 


/ / 




/ / 




' ' ,* 


Ctenoplusia 


' ' / 






Diachrysia 




Thysanoplusia 
Anadevidia 




Polychrysia 


/ '/s^ / 




'Jy^ / >• 


Euchalcia 


#y^ ________/ • 




f^ N. S 






Plusidia 




Lamprotes 


\ ,• 


Erythroplusia 


\ / 




\ / r 


Macdunnoughia 


\ / / 




\ s / 






Autographa 



Plusia 



Syngrapha 

Fig. 2 Phylogenetic tree of Ichinose. This tree originally consisted of a series of interconnected circles 
representing the genera. Thus several taxa (Abrostola, Chrysodeixis, Ctenoplusia (as Chrysodeixis) 
agnata, Euchalcia, Thysanoplusia (as Diachrysia), Erythroplusia and Autographa (s.str.)) were inter- 
preted as ancestral to other genera. These have been placed in the above figure on the ends of 
'zero-length' branches, represented by dashed lines. Only two character complexes were included on 
Ichinose's tree; larval chaetotaxy type and the presence/absence of lateral tufts on the adult male 
abdomen. Generic names have been changed to their modern forms. (Redrawn from Ichinose, 1962b). 



SYSTEMATICS OF THE PLUSIINAE 83 

asymmetrical male genitalia, larval morphology and, tentatively, larval foodplant. However, 
some possible affinity between Antoculeora and Macdunnoughia was admitted on the basis of 
the female genitalia (the sole species of Antoculeora, ornatissima , had been previously included 
by Ichinose [1962b] in Macdunnoughia) . Ichinose (1973) remained committed to the view that 
the Chrysodeixis-Trichoplusia-Argyrogramma complex formed a homogeneous group. To 
recognize a multitude of genera, as had Dufay (1970a), was regarded as 'myopic splitting'. 
However, Ichinose did relent somewhat and accepted Chrysodeixis (with Pseudoplusia as a 
tentative synonym) as distinct from Argyrogramma. This latter genus was divided into three 
subgenera: Argyrogramma, Trichoplusia and Ctenoplusia. Agrapha [Allagrapha] was synony- 
mized again with Plusia [Diachrysia] because the two shared many similarities in the form of 
abdominal segment 8 in the adult female and in the larval chaetotaxy (but see Eichlin & 
Cunningham, 1978, and below). 

At this point, I will digress briefly from the history of the higher classification of the Plusiinae 
to examine the problems associated with the type species of the genera Plusia and Diachrysia, 
and the application of these names. Up until 1970, most authors had treated orichalcea sensu 
Hubner, designated by Dyar (1902), as the type species of Diachrysia. This proved to be a 
misidentification of Diachrysia chryson and resulted in the orichalcea-intermixta group of 
species, which were certainly not congeneric with chryson (Kostrowicki, 1961 ; Ichinose, 19626) , 
being bereft of a genus. Ichinose (1973) resolved this by proposing Thysanoplusia. 

With regard to Diachrysia and Plusia, most authors had treated these genera as synonyms. In 
this they followed Grote (1895) in accepting DuponcheFs (1829) designation ofchrysitis as the 
type species of Plusia, a species that was then, and still is, considered to be congeneric with 
Diachrysia chryson. However, they had all overlooked DuponcheFs (1826) previous designa- 
tion of festucae as the type species. Thus Plusia correctly refers to the festucae-group of plusiines 
and Chrysaspidia sinks as a synonym. The next available name for the chrysitis-chryson group is 
Diachrysia. However, as noted above (and by Dufay, 1970a, and Nye, 1975), this name involves 
a case of misidentified type species, which should therefore be referred to the International 
Commission on Zoological Nomenclature under Article 70(b) of the Code. Although this will be 
done in the future, the consensus of all modern works (e.g. Franclemont & Todd, 1983; 
Bretherton et ai, 1984; Skinner, 1984) is that chryson is the type species of Diachrysia. 
However, the erroneous inclusion of orichalcea in Diachrysia is still prevalent (e.g. Bretherton 
etal, 1984; Skinner, 1984). 

To return to the higher classification of the Plusiinae, a totally independent test of all previous 
arrangements was performed by Nakamura (1974), using characters derived from the hitherto 
generally ignored pupal stage. Mosher (1916) and Gardner (19486) had examined a small 
number of species but Nakamura studied 21 species, which allowed him to comment on possible 
generic interrelationships. Very few of the groupings that he found corresponded to those 
previously proposed and several cut across the generic boundaries of Kostrowicki, Dufay and 
Ichinose. Congeners in Chrysodeixis (s.l. or s.str.), Thysanoplusia and Acanthoplusia showed 
marked differences in pupal chaetotaxy, while, in contrast, species of the genera Abrostola, 
Polychrysia, Plusia [Diachrysia] and Autographa (s.str., sensu Ichinose, 19626) had discrete 
pupal types, in terms of structure and chaetotaxy. Nakamura compared his results with the 
generic concepts of Ichinose (19626), Dufay (1970a) and Esaki et al. (1971) but refrained from 
making any formal taxonomic changes of his own. 

However, the latest Japanese checklist (Sugi, 1982) has returned to an almost maximally split 
arrangement. Lamprotes, Macdunnoughia, Antoculeora, Sclerogenia, Trichoplusia. Ctenoplu- 
sia and Acanthoplusia were all treated as distinct genera. Only Thysanoplusia (N.B.) and 
Puriplusia were recognized as subjective synonyms, being sunk to Trichoplusia and Macdun- 
noughia respectively. 

The focus moves back west: Eichlin & Cunningham's reappraisal of the Nearctic fauna 

Meanwhile, in the U.S.A. , analysis of the American Plusiinae had proceeded along similar lines 
as had the Japanese fauna a decade earlier. A series of studies of larvae (Eichlin & Cunningham, 



84 



IAN J. KITCHING 



1969; Eichlin, 1975) led eventually to a full scale revision of the subfamily on a regional basis 
(Eichlin & Cunningham, 1978). Like Ichinose, Eichlin & Cunningham employed both larval 
and adult characters in the construction of their classification. However, they tried not to place 
undue emphasis on either data source and so hoped to arrive at an arrangement that was less 
conservative than that of Ichinose but not as split as those of McDunnough, Kostrowicki or 
Dufay. The end result, however, did tend more towards the classifications of the latter three 
authors (Fig. 3). 

Largely on the basis of the form of the larval prolegs, Eichlin & Cunningham divided the 
subfamily into three tribes, the first occasion on which this taxonomic rank had been used in the 
Plusiinae. The most primitive tribe, in which the prolegs on A3 and A4 (see Terminology) were 
fully functional, if albeit slightly reduced, was the Abrostolini, comprising Abrostola and 
Mouralia. The latter was seen as a derivative of the former, having more specialized genitalia. 

Although Eichlin & Cunningham appeared to treat the Abrostolini as a monophyletic unit, 
they considered that 'a common ancestor from the tinctoides [as opposed to the Abrostolini] line 
gave rise to the Argyrogrammini [Argyrogrammatini] and Autographini [Plusiini]'. In both of 
these tribes, the ribs on the larval mandibles bore subterminal processes and the prolegs on A3 
and A4 were non-functional. 

The Argyrogrammini [Argyrogrammatini] were interpreted as the more primitive tribe 
because its members retained vestigial prolegs whereas all traces of prolegs had been lost in the 
Autographini [Plusiini]. Eichlin & Cunningham retained the three genera of McDunnough 




Syngrapha 
Plusia 



Fig. 3 Generic phylogeny of the Plusiinae according to Eichlin & Cunningham. Their original dendro- 
gram included an anagenetic component that is omitted in the above figure. The positions of Eosphoro- 
pteryx and Chrysanympha were uncertain, as indicated by dashed lines. Autographa was interpreted as 
being biphyletic. The detailed interrelationships of the species of Autographa and Syngrapha are 
omitted. (Redrawn from Eichlin & Cunningham, 1978.) 



SYSTEMATICS OF THE PLUSIINAE 85 

(Argyrogramma, Trichoplusia and Pseudoplusia) , rejecting Ichinose's (1973) synonymy of the 
first two, and of the third with Chrysodeixis, on the grounds that Ichinose's larval characters 
broke down when applied to the Nearctic species. Eichlin & Cunningham preferred to steer a 
middle course between Ichinose's larval dominated generic concepts and Dufay's adult domin- 
ated system, recognizing, in addition, that the results of Nakamura's (1974) pupal study were at 
odds with both. However, there were too few Nearctic Argyrogrammini [Argyrogrammatini] to 
permit Eichlin & Cunningham to draw any significant conclusions regarding the generic limits 
and interrelationships within the tribe. The three genera were represented on the phylogenetic 
tree (Fig. 3) as a trichotomy, although Pseudoplusia was considered to be the most advanced 
genus because it had lost the lateral abdominal tufts in the male (present in Trichoplusia and 
Argyrogramma), retained the mandibular processes (lost in the other two genera) and had a 
unique highly-elongate apex to the corpus bursae. 

The third tribe of plusiines, the Autographini [Plusiini], contained the balance of the 
American genera, 12 in all. Eichlin & Cunningham refrained from subdividing the group using 
adult characters because of the high degree of larval uniformity. The first group of four genera to 
branch off the main phylogenetic line were characterized by the retention of separate pinacula 
for setae SV1 andSV2on A2. In all other genera of Autographini [Plusiini] examined, these two 
pinacula were at least partially fused. Using the degree of separation of the VI setae on Al-4 
(supposedly linked to the time elapsed since the prolegs were lost on A3-4), Autoplusia was 
interpreted as the most primitive genus, followed by Rachiplusia, Plusia [Diachrysia] and 
Allagrapha. However, this arrangement was complicated by the presence of tibial spines, long 
believed to be an advanced character, in the former two genera. 

With regard to Allagrapha, Eichlin & Cunningham rejected its synonymy with Plusia 
[Diachrysia] . In particular, the presence of a raduloid (a ridged structure on the larval 
hypopharynx) in the two species of Allagrapha (absent in the Nearctic Plusia [Diachrysia] 
balluca Geyer, the only species of the genus examined as a larva), features of the male genitalia 
and a strong superficial similarity were used to argue full generic status for Allagrapha. 

The next section of Eichlin & Cunningham's tree was highly tentative because the larvae of 
four of the five genera (Adeva, Pseudeva, Eosphoropteryx and Chrysanympha)were not known. 
The fifth, Polychrysia, was considered to form the next branch off the main stem, because the 
larva retained the primitive arrangement of the V setae mentioned above. Adeva and Pseudeva 
were placed together because of the very reduced clavus and associated with Polychrysia 
because of the overall similarity of their genitalia. Chrysanympha and Eosphoropteryx were 
inserted next on the tree but their exact relationships were left open. 

Eichlin & Cunningham noted that Autographa, while structurally uniform in the male and 
female genitalia, could be subdivided into two lineages by the presence or absence of seta SV2 
on Al, presence being the derived condition. The final two genera, Syngrapha and Chrysaspidia 
[Plusia], were well defined. The former had spinose tibiae and the latter had uniordinal 
crotchets (unique in the Plusiinae) and an unusual larval mandible. Syngrapha was further 
subdivided into a number of species groups, one of which (including Palaeographa and 
Caloplusia) was characterized by the presence of seta SV2 on Til and Till, a feature unique 
within the Noctuidae (see also Crumb, 1956). 

Eichlin & Cunningham concluded by noting that the three major classifications, their own and 
those of Kostrowicki and Ichinose, were not particularly different in terms of the major 
evolutionary lineages they depicted. They considered this to be highly significant, given the 
more or less exclusive nature of the faunas upon which each was based. Nevertheless, Eichlin & 
Cunningham felt that there was still a great gap in the knowledge of the tropical plusiines and 
that this needed to be filled before a synoptic classification of the subfamily could be established. 

The latest North American checklist (Franclemont & Todd, 1983) explicitly followed Eichlin 
& Cunningham's tribal and generic assignments but within a few (unexplained) changes. 
Trichoplusia oxygramma was removed into Agrapha, under which was synonymized Ctenoplu- 
sia and Acanthoplusia. Allagrapha aereoides Grote (as aeroides) was put back into Diachrysia 
(following McDunnough, 1944) and Adeva was synonymized with Euchalcia (following Kostro- 
wicki, 1961). Anagrapha was resurrected for the divergent falcifera. 



86 IAN J. KITCHING 

Return to the east: Chou & Lu's phytogeny 

The late 1970s saw a surge of interest in the Plusiinae of China. Following a series of descriptive 
papers (Chou & Lu, 1974; 1978; 1979a; b), which included many inadvertent synonyms due 
probably to poor access to previous publications on the subfamily, Chou & Lu (1979c) published 
a summary of their ideas on the higher classification of the Plusiinae (Fig. 4). 




Fig. 4 Phylogeny of the Plusiinae according to Chou & Lu. Seven tribes were recognized (Omorphinini, 
Abrostolini, Euchalciini, Plusiini [sic], Caloplusiini, Argyrogrammini [sic] and Autographini) corres- 
ponding to the seven major lineages. Neoplusia and Scleroplusia are synonyms of Chrysodeixis and 
Macdunnoughia respectively and are placed in square brackets. Generic names have been converted 
into their modern forms. (Redrawn from Chou & Lu, 1919b.) 



SYSTEMATICS OF THE PLUSIINAE 87 

Seven tribes were recognized. The most primitive was considered to be the Omorphinini, 
comprising only Omorphina aurantiaca. Chou & Lu acknowledged the earlier suggestion 
(Kostrowicki, 1961) that Omorphina ought probably be transferred to the Erastriinae [Acon- 
tiinae], but they chose to retain it in the Plusiinae pending more detailed study. The next tribe, 
the Abrostolini, contained only Abrostola. Euchalcia, Chrysoptera [Lamprotes], Plusidia, 
Polychrysia and Panchrysia were placed in the Euchalciini, while the [Plusiini] comprised Plusia 
[Diachrysia], Thysanoplusia and Anadevidia. The Caloplusiini included Syngrapha and Calo- 
plusia. By far the largest tribe was the Argyrogrammini [Argyrogrammatini], consisting of four 
subgroups; two contained only single genera (Trichoplusia, Dactyloplusia), a third included 
Neoplusia [Chrysodeixis], Shensiplusia and Chrysodeixis, while the fourth consisted of a 
pentachotomy subtending Zonoplusia, Argyrogramma, Acanthoplusia, Puriplusia and Erythro- 
plasia. These last two genera were placed near Acanthoplusia despite lacking vestigial larval 
prolegs (also absent in A. agnata; Ichinose, 1962b) and anal tufts in the male. If Puriplusia and 
Erythroplusia are excluded, then the Argyrogrammini [Argyrogrammatini] is more or less 
equivalent to Chrysodeixis (s.l.) (sensu Ichinose, 1973). 

The final tribe, the Autographini [Plusiini] , included six genera in three subgroups. Chrysaspi- 
dia [Plusia] represented the most basal branch, followed by Autocureola [sic, Antoculeora]. The 
four remaining genera (Autographa, Cornutiplusia, Scleroplusia [Macdunnoughia] and Mac- 
dunnoughia) were treated as a tetrachotomy. Cornutiplusia was considered to be identical to 
Autographa (except for the spined tibiae) and Autocureola [Antoculeora] was interpreted as the 
most advanced genus. 

No resolution of the tribal interrelationships was attempted. Several of the tribes, notably the 
[Plusiini], were distinctly phenetic in construction. Chou & Lu preferred to group genera on a 
limited number of superficial characters of doubtful taxonomic significance (see below), which 
resulted in a number of generic misplacements. Nevertheless, the overall classification had a 
great deal in common with those proposed previously. 

Ronkay (1982), in a review of the Korean fauna, also accepted the general findings of 
Kostrowicki, Ichinose and Eichlin & Cunningham. However, he disagreed on several points. 
Ronkay considered Ichinose's interpretation of Macdunnoughia as a subgenus of. Autographa 
was incorrect and he reinstated the former to generic level. Chou & Lu's ( 1974; 1979c) treatment 
of Puriplusia as a good genus was also disputed and Ronkay, agreeing with Dufay (1977), 
reduced Puriplusia to subgeneric rank within Macdunnoughia. The generic assignments within 
the Argyrogrammatini were also considered problematical and, pending a full tribal revision, 
Ronkay chose to retain the genera Trichoplusia, Ctenoplusia, Chrysodeixis and Argyrogramma 
according to Dufay's usage. 

Holloway (1985), in his study of the Bornean fauna, also maintained Chrysodeixis and 
Ctenoplusia as distinct, pending further investigation. Thysanoplusia, Zonoplusia and Dactylo- 
plusia were also treated as good genera but several plusiine species remained unassignable to 
current genera. Holloway followed the practice of Dufay (e.g. 1972; 1974) in placing such 
species in 'Plusia' s.l. Although this approach is unsatisfactory, it was the best that could be 
achieved for these species until a comprehensive revision of the subfamily, and particularly the 
Argyrogrammatini, is performed. 

Kljutshko's 'cladistic' analysis of the Plusiinae 

The most recent studies to consider the higher classification of the Plusiinae are those of 
Kljutshko (1985a; b). For the first time in plusiine systematics, cladistic techniques were overtly 
employed to analyse a data set of 32 characters, drawn from larval, pupal and adult morphology. 
The result was a cladogram (Fig. 5) based upon the 24 characters that Kljutshko considered to be 
the most well-founded. However, the analysis was fundamentally flawed in such ways as to 
render most of the resolution of the cladogram spurious and thus invalidate most of the 
conclusions drawn from it. 

First, and most crucial, the cladistic methodology was misapplied. Although some groups 
were supported by putative synapomorphies (e.g. Ctenoplusia and Acanthoplusia on the basis of 



88 



IAN J. KITCHING 



Autographa 

Desertoplusia 

Diachrysia 

Macdunnoughia 

Antoculeora 

Plusia 

Plusidia 

Euchalcia 

Pseudochalcia 

Anadevidia 

Lamprotes 

Panchrysia 

Polychrysia 

Cornutiplusia 

Syngrapha 

Caloplusia 

Trichoplusia 

Argyrogramma 

Chrysodeixis 

Zonoplusia 

Ctenoplusia 

Acanthoplusia 

Erythroplasia 

Abrostola 

Fig. 5 Cladogram of the plusiine genera according to Kljutshko. (Redrawn from Kljutshko, 1985a, b.) 

blade-like setae on the valve), others were distinguished and delimited using symplesiomor- 
phies. For example, Autographa and Desertoplusia were united, and separated from Diachrysia 
on the basis of a long harpe, the plesiomorphic state for his character 22. Similarly, the members 
of the 'clade' Autographa- Antoculeora were distinguished from Plusia by the presence of 
biordinal crotchets on the larval prolegs (Kljutshko's character 6). However, all other known 
plusiine larvae have biordinal crotchets and it is more parsimonious (and in keeping with 
previous authors' conclusions) to treat the uniordinal condition in Plusia as autapomorphic for 
this genus. This would appear to be a failure to employ the outgroup criterion correctly. 

If character 6 is reinterpreted and all the groups characterized by symplesiomorphies 
collapsed to form polytomies, then a much less well-resolved cladogram results (Fig. 6). Three 
tribes were recognized by Kljutshko: the Abrostolini, the Argyrogrammatini (including 
Erythroplusia) and the Plusiini. The last consisted of three subtribes: the Plusiina (including 
Anadevidia), the Caloplusiina and the Polychrysiina. Of the tribes, the Abrostolini and the 
Argyrogrammatini were sister-groups, in contrast to every other phylogeny that had been 
previously proposed for the Plusiinae. This relationship was based upon the possession of a 




SYSTEMATICS OF THE PLUSIINAE 



89 



Autographa 

Desertoplusia 

Diachrysia 

Macdunnoughia 

Antoculeora 

Plusia 

Plusidia 

Euchalcia 

Pseudochalcia 

Anadevidia 

Lamprotes 

Panchrysia 

Polychrysia 

Cornutiplusia 

Syngrapha 

Caloplusia 

Trichoplusia 

Argyrogramma 

Chrysodeixis 

Zonoplusia 

Ctenoplusia 

Acanthoplusia 

Erythroplusia 

Abrostola 

Fig. 6 Cladogram of the Plusiinae derived from the data set of Kljutshko (19856) following the correct 
application of cladistic methodology. Many of the taxa in Fig. 5 were delimited by plesiomorphic 
character states. Their elimination results in numerous polytomies. 




finer, hair-like seta SD1 on A9 of the larvae (this state is also convergently derived in 
Cornutiplusia). 

However, even the reduced cladogram shown in Fig. 6 is defective. The reason for this lies in 
the characters that Kljutshko used, many of which are insubstantial and nugatory (e.g. the forms 
of the clavus and harpe, and the origin of the ductus bursae on the corpus bursae), and all of 
which appear to have been culled from the works of other authors. The result suffers from 
deficiencies similar to those in the 'phylogenetic tree' of Chou & Lu (1979c). 

Within the constraints imposed by Kljutshko's choice of characters, the following additional 
modifications can be made to the character analysis (albeit in the light of my own analysis 
described below). In addition to character 6, the polarity of character 26 should be reversed 
because the absence of a signum is plesiomorphic for the Plusiinae as a whole. Since the absence 
of an epicranial suture in the pupa characterizes the members of both the Plusiini and the 
Argyrogrammatini (see also Nakamura, 1974), this state can be reinterpreted as synapomorphic 



90 



IAN J. KITCHING 



for these two tribes. It also follows from this that the polarity of character 4 (SD1 on larval A9) 
should be reversed as well. I also reject characters 12 (only Euchalcia gerda and E. serraticornis 
Dufay [Dufay, 1968] have serrate antennae, and only Lophoplusia has bipectinate antennae, 
none of which were examined in Kljutshko's study), 18 (if the dorsal abdominal tuft on the adult 
A2 is reduced in any genus, it is in Plusia, not Syngrapha; see almost any author on the Plusiinae) 
and 29 (the position of the ostium bursae on A8) as having any significance. The final result of 
these changes is shown in Fig. 7 and represents the only informative structure present in 
Kljutshko's original cladogram (Fig. 5). 

As alluded to above, the Argyrogrammatini and Plusiini have a sister-group relationship but a 
trichotomous one that includes Erythroplasia as an unplaced genus. The members of the 
Argyrogrammatini are united by the presence of subdorsal tufts on A5-6 in the male but form an 

Autographa 

Desertoplusia 

Diachrysia 

Macdunnoughia 

Antoculeora 

Plusia 

Plusidia 

Euchalcia 

Pseudochalcia 

Anadevidia 

Lamprotes 

Panchrysia 

Polychrysia 

Cornutiplusia 

Syngrapha 

Caloplusia 

Trichoplusia 

Argyrogramma 

Chrysodeixis 

Zonoplusia 

Ctenoplusia 

Acanthoplusia 

Erythroplasia 

Abrostola 

Fig. 7 Cladogram of the Plusiinae derived from the data set of Kljutshko (1985b) following further 
character analysis. Several characters (see text) were reinterpreted by the correct application of the 
outgroup criterion for determining character polarity and several more were omitted. The resulting 
cladogram represents the sum total of the information in Kljutshko's data set. There are extensive 
polytomies and few informative groups. Note, however, the monophyly of the Argyrogrammitini, 
Ctenoplusia (s.l.) and the clade Macdunnoughia + Antoculeora. 




SYSTEMATICS OF THE PLUSIINAE 91 

apical pentachotomy, in which the only further resolution is the union of Ctenoplusia and 
Acanthoplusia on the basis of blade-like setae on the valve. The Plusiini are characterized by a 
normal seta SD1 on A9 and the total loss of prolegs in the larvae. The 16 constituent genera of 
this tribe also form a highly polytomous apex, with no fewer than ten branches. Euchalcia and 
Pseudochalcia are united by the form of the juxta, while Macdunnoughia and Antoculeora share 
a weak uncus that is not apically hooked. Polychrysia, Lamprotes and Panchrysia are placed 
together by the possession of very long palps and probosces but form a trichotomy. Cornutiplu- 
sia, Syngrapha and Caloplusia are united by the shared presence of tibial spines, while the latter 
two genera are treated as sister-groups because they both display a juxta with an apical process 
(the same state that unites Euchalcia and Pseudochalcia). All the genera have autapomorphies 
that will not be considered further here, with the exception of Chrysodeixis, Ctenoplusia, 
Autographa, Polychrysia, Lamprotes and Syngrapha. 

I have conducted the preceding re-analysis of Kljutshko's cladogram in order, not to deride 
his work, but to congratulate him. He, amongst all previous authors (with the possible exception 
of Ichinose, 19626), has produced a thoroughly explicit analysis that is open to examination and 
constructive criticism. Many other authors (notably Kostrowicki [1961] andChou& Lu [1979c]) 
chose not to state the exact reasons (if any) for the conclusions they reached. Even Eichlin & 
Cunningham ( 1978) only explicitly stated some of the characters they used in order to reach their 
final classification, which rendered many parts of their 'phylogenetic tree' immune to criticism. 
Thus, despite the errors in analytical procedure and his reliance upon a small number of largely 
superficial characters (see below), Kljutshko did produce an arrangement for which his 
reasoning was clear and in which the characters used to delimit each taxon were unambiguously 
stated. 

Two unconventional approaches - 

All the studies discussed above investigated the external morphology of the larvae, pupae and 
adults, with primary attention being focussed on the cuticular structures of the male and female 
genitalia. There have been two further studies that considered more unconventional organ 
systems. Both deal with only a very restricted sample of species but the results do appear to 
corroborate some aspects of previous classifications. For this reason, they will be discussed in 
detail. 

Callahan (1960) examined the structure of the male and female reproductive tracts in eight 
species of Plusiinae, paying particular attention to those parts that operate in the formation and 
insertion of the spermatophore. Thus, in addition to the aedeagus and the corpus bursae, 
Callahan also examined the cuticular simplex (the part of the male genitalia between the testes 
and the aedeagus that is normally removed in genitalia preparations) and the spermatophore 
itself. 

The species were found to fall into two distinct groups. Group I included Autographa biloba, 
Agrapha [Allagrapha] aerea and Rachiplusia ou (Guenee), and thus contained only and all the 
members of the Plusiini studied. They were characterized by a spermatophore with a short, 
heavy collum and large cap, a short aedeagus without a sclerotized bulbous lock and a cuticular 
simplex with a short, broad constrictor muscle. In contrast, the species of Group II had a 
medium or long, narrow collum, a small spermatophore cap that fits tightly at the apex of the 
corpus bursae where the ductus seminalis arises, a sclerotized bulbous lock on the inner wall of 
the uneverted vesica, a long aedeagus and a constrictor muscle that is elongate and tapered at the 
basal end. This group included Pseudoplusiaincludens, Trichoplusiani, T. oxy gramma (Gey er) , 
Argyrogramma verruca and A. basigera (Walker), and thus corresponded to the Argyrogram- 
matini. Although there were characters that might have allowed subdivision of these groups 
(particularly states that might group Autographa and Agrapha [Allagrapha] to the exclusion of 
Rachiplusia; and Trichoplusia and Pseudoplusia to the exclusion of Argyrogramma), such 
analysis was precluded by the lack of an outgroup by which to polarize the transformation series. 
However, the results of Callahan's study did corroborate the tribes subsequently proposed by 
Eichlin & Cunningham (1978). 



92 IAN J. KITCHING 

The second 'unconventional' morphological study examined the musculature of the male 
genitalia (Bayer, 1965). The results of a previous (unreferenced) study of the valves of 25 South 
African plusiine species suggested that they could be divided into five groups. Four (A-D) 
corresponded to Cornutiplusia, Chrysodeixis, Ctenoplusia and Trichoplusia (part). The fifth 
group, E, included two species of Thysanoplusia, two species of Trichoplusia (sestertia (Felder) 
and arachnoides (Distant)) and 'Plusia' tetrastigma (Hampson) (which may properly belong in 
Trichoplusia [Dufay, note in BMNH collection]). Finally, there were four unplaced species: two 
Abrostola and two Trichoplusia (transfixa (Walker) [vittata (Wallengren)] and obtusisigna 
(Walker)). The distinction between groups D and E was noted as not being very marked. Bayer 
then proceeded to examine the male genital musculature of six species in greater detail: 
Cornutiplusia circumflexa, Chrysodeixis acuta (Walker), Ctenoplusia limhirena, Trichoplusia 
angulum (Guenee), T. transfixa [vittata] and Thysanoplusia orichalcea. (The first species was 
placed in Syngrapha, the remainder in Plusia.) 

Although a number of structural differences were found to suggest Cornutiplusia, Chry- 
sodeixis and Trichoplusia vittata might be separated from the other three species, no characters 
were found that would enable their grouping into higher taxa. Nevertheless, the discovery of 
quite major musculature changes, such as the loss of entire muscles or marked differences in 
insertions, does suggest that the genital musculature has great potential in higher classification. 

The position of the Plusiinae within the Noctuidae 

When the Plusiinae were first proposed as a group (Boisduval, 1828), they were placed as part of 
a linear sequence between the Noctuelidi, a conglomerate of the presently recognized trifine 
subfamilies, and the Heliothidi. Guenee (1841) and Duponchel (1844) retained a tentative 
relationship with the Heliothidi, but also favoured an affinity with the Calpidi. In contrast, 
Herrich-Schaffer (1845) removed the Plusidae [sic] from any association with the Heliothidae, 
and placed them in a part of his sequence that included the Metoponidae and Erastridae. 

Guenee's second classification (1852-4) divided the noctuids into two main phalanges based 
on hindwing venation. The Plusidae were included in the tribe Variegatae of the phalange 
Quadrifidae, together with such families as the Eurhipidae [Euteliinae], Calpidae and Gonop- 
teridae. Thus, in his revised classification, Guenee retained a relationship between the Plusidae 
and the Calpidae, but rejected a connection with the Heliothidae, which was placed in the 
phalange Trifidae. 

Packard (1869) also employed a two-way split of the Noctuidae but based upon the shape of 
the clypeus and the antennal form, rather than wing venation and palps. Thus, he included the 
plusiines in the Noctuinae with the trifine noctuids, rather than in the Catocalinae, with the 
quadrifines. Grote (1882; 1895) continued this usage, which reassociated the Plusiinae with the 
Heliothinae, but also placed the Calpinae nearby, together with the Stiriinae, which were 
considered to be closely related to the heliothines. 

Both Tutt (1891-2) and Hampson (1893-5) preferred to use versions of Guenee's (1852-4) 
second classification. Tutt (1902) subsequently modified his position to approximate that of 
Grote (1895) by considering the plusiines to be a highly developed group of heliothines. The 
reduced larval prolegs of most plusiines, by which they were associated with the Calpinae and 
other quadrifines with semi-looper larvae, were explained as the result of convergent evolution. 

In contrast, Hampson (1902) had completely reappraised his classification of the Noctuidae 
and this culminated in a system that was at once to revolutionize and stultify noctuid higher 
systematics. By classifying the noctuid moths into semi-artificial assemblages, using a small 
number of characters applied absolutely, Hampson succeeded in creating a series of subfamilies 
between which there were no possibilities for higher level relationships. Noctuid higher 
classification, which had been hierarchical to varying degrees since Boisduval (1828), had been 
reduced to a single tier system in one fell swoop. The Plusiinae were no longer seen as having 
closer relationships with some subfamilies than others; now they were just one among 15 equals. 
Even the major division between trifine and quadrifine hindwing venation was not reflected in 
the classification. 



SYSTEMATICS OF THE PLUSIINAE 93 

This attitude towards higher level classification was further institutionalized by the authors of 
'Seitz\ but this unfortunate state of affairs did not long go unchallenged. Richards (1932) 
conducted a detailed comparison of the morphology and evolution of the tympanal organ in 
which, in addition to dividing the Noctuidae into a number of subfamilies, he also suggested 
possible interrelationships between the subfamilies in the form of a phylogenetic tree. The 
Plusiinae were found to exhibit a very distinct type of tympanal organ, with a double tympanal 
hood and a greatly enlarged pocket IV, intimately fused to the tympanal air sac. The subfamily 
Stictopterinae were found to possess a tympanal hood with a ventral lobe and a pocket IV which, 
while of normal size for a noctuid, was intimately fused with the air sac as in the Plusiinae. On the 
basis of these similarities, Richards proposed that the Plusiinae had been derived via the 
Stictopterinae from the higher Erebinae [Catocalinae] of group IV (including Euparthenos 
Grote and Zale Hiibner). This relationship between the Stictopterinae and the Plusiinae was 
accepted by all subsequent authors, Forbes (1954) even going so far as to suggest that the two 
subfamilies might better be treated as one. Eichlin & Cunningham (1978) also found no fault 
with this suggestion and used it in order to construct their 'prototype' plusiine. One of the major 
conclusions of the work detailed below is that the belief in such a stictopterine-plusiine 
sister-group relationship is erroneous. 

Taxa examined 

The 400 or so species of Plusiinae are currently classified into about 30 to 50 genera, depending 
upon the degree of taxonomic 'splitting' adopted. To attempt to maximise the resolution of the 
supraspecific relationships and to test the validity of many of the synonymies, 45 nominal genera 
were selected for analysis, all of which have been in recent use. The type species of each 
(Appendix 1) was employed as an exemplar taxon and although difficulties were encountered in 
coding some characters that displayed intrageneric variation, this approach was adopted to 
minimise ambiguity in the application of generic names. In addition, two other plusiine taxa 
were included: Palaeographa Kljutshko, which had been proposed only as a subgenus of 
Syngrapha Hiibner; and Autographa bractea ([Denis & Schiffermuller]), to represent the more 
derived lineage of Autographa Hiibner, a genus considered to be biphyletic by Eichlin & 
Cunningham (1978) (but see Clade analysis). 

Several plusiine generic level taxa were omitted. These are given in Appendix 2, together with 
the reasons for their rejection. Unjustified emendations and subsequent misspellings are not 
included. Several additional plusiines, other than the type species, were dissected for a variety of 
reasons. 

Antoculeora locuples (Oberthur, 1881): the BMNH series of Antoculeora ornatissima may include one or 
more of this taxon, which were dissected in error. However, this did not affect the present study for the 
putative A. locuples specimen (1 d") did not differ from the putative A. ornatissima in any character 
scored in this study. 

Autoplusia olivacea (Skinner, 1917) (Fig. 81): dissected by chance from a mixed series with A. egena but 
which proved useful in confirming several character states and transformation series. 

'Syngrapha' gammoides (Blanchard, 1852) (Figs 82, 83): dissected for another study and used in the 
present investigation because the male genitalia were less specialized than those of either A. egena or A. 
olivacea and thus aided the clarification of several characters. '5. ' gammoides is almost certainly a species 
of Autoplusia. This genus is currently being investigated by Fernando Navarro (Tucuman) and pending 
the results of his study, I will refer to this moth as 'Syngrapha gammoides. 

Chrysodeixis argentifera (Guenee, 1852) (Fig. 40); C. illuminata (Robinson, 1968) (Fig. 41); C. kebea 
(Bethune-Baker, 1906) (Fig. 42): the type species of Chrysodeixis, chalcites, has extremely modified 
sternal hair pencils in the male (see Terminology) and it proved impossible to score characters 80-95 for 
this species. Three further species of Chrysodeixis were dissected in order to determine the nature of the 
modification in C. chalcites. C. kebea and C. illuminata were found to be almost as extreme as C. chalcites 
but C. argentifera possessed a typical pair of sternal hair pencils. Characters 80-95 were thus scored from 
this latter species. 

Plusiopalpa shisa Strand, 1919: only a single male of the type species, P. dichora, was found in the BMNH 
collection, and so a male of P. shisa was also dissected to check leg and abdominal characters. The 



94 IAN J. KITCHING 

identification of this specimen as P. shisa rested solely on its Taiwanese origin. A study of the genitalia 
may demonstrate this species to be synonymous with P. adrasta (Felder). 

In addition, 11 outgroup taxa were analysed (Appendix 1) and were chosen on the following 
bases. 

Stictopteral Lophoptera: to represent the two main lineages (Holloway, 1985) of the subfamily Stictopter- 
inae, which is considered by many to be closely related to the Plusiinae. The BMNH series of the type 
species of Lophoptera, L. squammigera , appears to be mixed (J. D. Holloway, pers. comm.) but this will 
have no effect on the present study. 

Paectes/Anuga: to represent the subfamily Euteliinae, considered by Holloway (1985) to be a more likely 
candidate for the sister-group of the Stictopterinae. Paectes cristatrix was used inadvertently instead of 
the type species, P. pygmaea Hiibner. 

Magma: this genus superficially resembles Stictoptera and may be related to it. 

Calophasial Cucullia: to represent the subfamily Cuculliinae, which share with the Plusiinae the character 
state of lashed eyes. 

Diloba/ Brachionycha: Diloba has been associated in the past with the Plusiinae (amongst many other 
groups, see Kitching, 1984) and this relationship was to be tested. It has also been suggested that Diloba 
is related to the cuculliine tribe Psaphidini and thus Brachionycha was included, as the only member of 
this tribe sufficiently well represented in the BMNH collection to be utilized. 

ChalcopastalOncocnemis: the former genus has a similar facies to members of the plusiine genera 
Thysanoplusia and Diachrysia, and a relationship between the amphipyrine tribe Stiriini (of which 
Chalcopasta is a member) and the Plusiinae has been very tentatively suggested. However, Hogue 
(1963) , in his revision of the Stiriini, strongly favoured a relationship between this tribe and the cuculliine 
tribe Oncocnemidini, hence the inclusion of Oncocnemis as an outgroup in this study. 

Several other genera were initially examined but were rejected because they appeared to 
have no close affinity with the Plusiinae - Plusiophaes Prout, Scoliopteryx Germar, Xylena 
Ochsenheimer, Lithophane Hiibner, Nycteola Hiibner and Pseudoips Hiibner. In addition, two 
stictopterines, Odontodes Guenee and Nigramma Walker (now a synonym of Gyrtona Walker; 
Holloway, 1985) were scored but rejected as redundant to the problem at hand. 

All taxa were scored from specimens held in the BMNH collection except for the follow- 
ing: 

Loboplusia vanderweelei holotype d\ paratype cf: Rijksmuseum van Natuurlijke Historic Leiden, 

Netherlands; 
Autographa pokhara Dufay holotype $ (= Loboplusia vanderweelei $): Zoologische Staatssammlung, 

Munich, West Germany; 
Shensiplusia nigribursa holotype $: Northwestern College of Agriculture, Wukung, China. This species 

was not examined; however, as many characters as possible were scored from the original description 

and accompanying diagrams of wing pattern and female genitalia (Chou & Lu, 1974). 

Terminology 

In this section, I shall define those morphological terms that may have ambiguous usage within the 
Lepidoptera or that I have introduced in this study. They concern mainly internal structures, the male 
sternal hair pencils and the male genitalia. Unambiguous or universally accepted terms are omitted. 
Genitalic terminology generally follows Klots (1970); wing venation follows Neilsen (1985). Thoracic 
segments are termed TI— III (Roman numerals) and abdominal segments Al-8 (occasionally -A10 with 
reference to larvae). The abdominal tergites and sternites are termed Tl-8 (note Arabic numerals) and 
Stl-8 respectively. 

Wings. The scale tooth is a triangular tuft of scales projecting from the tornal angle of the forewing. In the 
males, the frenulum consists of a single bristle formed by the fusion of several setae. In most of the species 
examined in this study, the female frenulum consists of three setae. The innermost seta is termed 1, the 
outermost 3 (Fig. 124). Reduction in number generally results from the loss of seta 1 first, and seta 2 next. 
Whether reduction is by loss or by fusion is often difficult to assess. In those species with four frenular setae, 
it is unknown whether the extra seta represents a '0' or a '4'. Crossveins are identified by the main veins 
between which they run, labelled in lower case. Thus m 1 -m 2 is the cross vein between Mj and M 2 . The 
terms lower, middle and upper discocellular veins are avoided. 



SYSTEMATICS OF THE PLUSIINAE 95 

Labial palps. The apical 'pit organ' of segment 3 is here termed 'vom Rath's organ' (Rath, 1887), following 
Kristensen ( 1984). The sensilla contained in the pit have been thought to be chemoreceptors, thermorecep- 
tors or hygroreceptors (Chauvin & Faucheux, 1981), but have recently been shown to be sensitive to 
carbon dioxide (Bogner etal. , 1986). 

External thoracic structure. The tegulae (Figs 164-167) are interpreted as having two arms. The dorsal 
arm is broad and usually bluntly pointed or rounded, and protects the forewing base dorsally. The ventral 
arm is thin and covered in microspines apically. It curves out and under the wings, where its spinose apex 
may aid in holding the wings in position at rest. 

Legs. The basitarsal comb is a row (or rows) of specialized setae (Figs 178-181) on the inner subventral 
edge of the hindbasitarsus of many male argyrogrammatines. The function of these combs is still unknown, 
although their partial correlation with the dorsolateral hair tufts (see below) suggests that they may be used 
for grooming. They do not seem to be absolutely essential, however, for although all species with combs 
have tufts, not all species with tufts have combs. The genus with the most highly developed tufts of all, 
Chrysodeixis, lacks combs. 

In general, there are three discrete rows of setae along the ventral surface of the tarsi in plusiines. The 
central row is termed the V (ventral) row, the lateral rows SV (subventral). Additional SD (subdorsal) and 
D (dorsal) setae may also occur. 

Internal prothoracic structure. The internal structure of the ventral part of the prothorax of a typical 
plusiine is illustrated in Fig. 211. The ventral midline (discrimen) is inflected to form two large, curved 
processes that form the dorsal arms of the prothoracic furca. Basally, these are produced posteriorly to 
form a thin, apically bifurcate spinisternum (not shown). The dorsal furcal arms are produced subventrally 
into broad, rounded projections termed the subventral (SV) furcal arms. To these structures are attached 
several of the foreleg muscles. The apices of the dorsal arms are connected on each side to the dorsal edges 
of the epimera by a cuticular membrane. This structure may possess a dorso-ventral ridge running from the 
epimera to the SV furcal arms (Fig. 212). 

Male abdomen Al-7. Terminology for the trifine hair pencil components follows that of Birch (1972a; b). 
Dorso-lateral hair tufts occur in many male Argyrogrammatini. The tufts have greasy, matted, apparently 
secretory patches of scales at their bases and are assumed to be scent organs, presumably involved in 
courtship. Those of A7 are occasionally enclosed in cuticular folds of various types. All are easily removed 
during normal preparatory procedure for slide mounting and then often go undetected. 

Male A8 and sternal hair pencils. The modifications to A8, concerning the elaboration of the sternal hair 
pencils of male plusiines, are very complex and still incompletely understood. The plesiomorphic condition 
was presumably a simple quadrate T8 and St8. In many of the genera examined, including all of the 
plusiines and several of the outgroups, T8 was cruciform, or bilobate, in which the anterior arms had been 
reduced or lost. The shape of T8 is often species specific (Dufay, 1970a) but although much variation was 
found in this study, no patterns to the different forms could be discerned and thus no characters were 
derived from it. 

St8 appears first to have evolved a central, posterior, desclerotized area giving rise to a U-shaped sclerite 
in which the arms point posteriorly. Between these are developed a patch of secretory, setose scales. This 
desclerotized area is then invaginated as two pockets (more rarely one), in which are located the sternal 
hair pencils. These pockets are often fused, partially or totally, and the hairs themselves occasionally 
differentiated into several types in terms of length , curvature , colour and basal insertion. The development 
of the pockets bent the U-shaped St8 in the dorso-ventral plane, resulting in a thin strap supporting the 
edge of the pockets ventrally, termed the anterior arm of St8, and two thin bars projecting posteriorly 
above the opening of the pockets, the lateral arms of St8. The membrane between the lateral arms can be 
variously scaled. 

In the Argyrogrammatini, the lateral arms of St8 are further modified. Initially, they develop apical 
secondary hair brushes. In some genera, the lateral arms also develop sub-basal, exterior lobes, which 
represent the attachment points of the secondary hair brushes. The part of the lateral arm that is distal to 
this lobe is referred to as the dorsal arm of St8, as it is a morphologically recognizable subdivision of the 
lateral arm that lies dorsal to the hair pencils. In some argyrogrammatines, the dorsal arms bear flat plates 
of greasy, closely appressed scales. 

As alluded to above, many Chrysodeixis species have even more modified structures on A8. C. 
argentifera is unmodified and is thus interpreted as being a relatively plesiomorphic species in terms of 



96 IAN J. KITCHING 

these structures. In other species, however, and especially C. chalcites and C. kebea, the sternal hair pencils 
have been lost entirely and St8 modified into a compact sagittate sclerite (Fig. 250). The function of the 
sternal hair pencils has been taken over by the secondary hair brushes, which have moved ventrally and 
become retracted into a shallow, semi-circular pocket. T8 has also become highly modified into a U-shaped 
sclerite with thin arms. It also has attenuated lateral and anterior arms that enclose and support the pocket 
of the secondary hair brushes (Figs 251-253). In consequence, T8 in the 'higher' Chrysodeixis has become 
remarkably convergent in form upon St8 in other argyrogrammatines, except that its components are 
rotated through 180 degrees relative to those of St8. 

The sternal hair pencils and associated structures of Trichoplusia ni have been described in detail by 
Grant (1971) and their role in courtship was explained by Gothilf & Shorey (1976). 

Tegumen and uncus. In most species examined in this study, there occur, latero-apically on the arms of the 
tegumen, two small sclerites that project posteriorly and under the base of the uncus. These are termed the 
'latero-apical sclerites of the tegumen' or LASTs. They may represent the remains of the gnathos since they 
occur 'ventro-laterad of the articulation of the uncus and socii if present' (Tuxen, 1970). 

Vinculum and pleurite. In this study, most of the species examined lacked a saccus in the male genitalia as 
commonly understood. Instead, the arms of the vinculum converge to a point. In several genera, dorsal and 
ventral flanges of these arms are produced and fused to form saccus-like structures but are referred to by 
their appearance rather than as sacci. 

The pleurite (termed the paratergal sclerite by Tikhominov, 1979) is a small sclerite, often curved and 
flanged, that is found exteriorly and laterally at the point where the vinculum and tegumen arms meet. In a 
few genera, the pleurite was seen to be continuous basally with the outer surface of the vinculum and thus 
appears to be an extension of the latter. In many plusiines, the base of the pleurite has become sharply 
kinked and may have lost its connection with the vinculum. In a small number of taxa, the pleurite has 
become fused with the dorsal edge of the tegumen or may have been lost altogether. In several genera, the 
vinculum arms continue posterior to the origins of the pleurites and these extensions are termed the 
ventral, apical, vinculum bars (VAVBs). 

Valve. There is much confusion and ambiguity concerning the terminology of the valves and their 
constituent structures. In this study, the following terminology is adopted. The valves are the paired, 
presumably clasping organs of the male genitalia. The dorsal edge of the valve is the costa; the ventral, 
often recurved edge, the sacculus. At the base of the sacculus is produced a usually tubular appendage 
termed the clavus, while more distally, a similar structure is termed the harpe. The harpes are much more 
variable in form than the clavi. The transtillae are sclerotized bars, produced from the bases of the costae, 
that project inwards and support the anellar membrane dorsal to the aedeagus. The aedeagus is supported 
ventrally by the juxta. 

Aedeagus. The muscular tube arising from the base of the aedeagus, down which the spermatophore 
passes, is termed the cuticular simplex (Callahan, 1960). If present, the enlarged section of the aedeagus 
basal to the origin of the cuticular simplex, especially if bulbous, is termed the coecum. References in the 
character descriptions of structures on the vesica as LHS (left hand side) and RHS (right hand side) are 
from the moth's point of view. 

Female Al-7. All of the taxa examined in this study, in common with the majority of the rest of the 
Noctuoidea, possess a counter-tympanal hood. This takes the form of a rounded, quadrate, sclerotized flap 
produced from the pleural region of segment A2. It is concave on its inner surface and curves forward to 
protect the metathoracic tympanal organ. The Euteliinae, Stictopterinae and Plusiinae were observed to 
possess a second hood-like structure, antero-ventral of the counter-tympanal hood, which is often referred 
to in the literature as a 'secondary tympanal hood'. Close examination, however, shows that it is a flattened 
pouch, the two walls of which are separate (Figs 230-231), not a single, solid structure, as is the 
counter-tympanal hood. I will therefore refer to it as the 'pleural pouch'. 

The inner and outer surfaces of the pleural pouch are supported by two separate sclerotized bars of 
markedly different origins. A latero-apical production of the outer margin of the anterior apodeme of St2 
(Figs 230-231) projects antero-laterally and then recurves sharply postero-ventrally to support the inner 
surface of the pouch . In contrast , the outer surface of the pouch is supported by a ventro-anterior extension 
of the post-spiracular bar (Fig. 231). 

In most euteliines, stictopterines and plusiines examined, the pleural pouch is a large structure, 
projecting antero-ventrally below the counter-tympanal hood. However, in Polychrysia and its relatives 



SYSTEMATICS OF THE PLUSIINAE 97 

(see character 178), the pouch is small and has moved dorsally so as to be partially concealed by the 
counter-tympanal hood (Fig. 232). 

Female genitalia. The correct homologies of segment 8 of the female noctuid have yet to be fully 
elucidated. For the purposes of this work, T8 will refer to the sclerotized semi-annulus encircling A8 
dorsally and laterally. The often membranous section between the edges of T8 ventrally is termed the 
lamella postvaginalis (LPV). The antrum is the initial differentiated section of the ductus bursae 
immediately internal to the ostium bursae. It is frequently more heavily sclerotized and usually much 
expanded relative to the rest of the ductus bursae. In general, the apex of the corpus bursae is the 
posteriorly-directed section posterior to the entry point of the ductus bursae into the corpus bursae. The 
fundus is the opposite , anterior end of the corpus bursae (but see also character 209) . 

Methods 

For each species (except Plusiotricha, Eutheiaplusia, Stigmoplusia, Chrysanympha, Adeva and Loboplu- 
sia), two specimens, one male and one female, were selected. First, general facies and pattern characters 
were scored from the intact animals. Then, the right fore and hindwings were detached, bleached and 
mounted in Euparal. From these, the wing structure characters were scored. Next, the head, prothorax, 
patagia, tegulae and legs were removed, heated in 10% KOH solution, descaled and, except for the head 
capsule and prothorax, mounted in Euparal. Most characters from these parts were scored before 
mounting but several required confirmation by observation using a phase contrast compound microscope 
at x400. Finally, the abdomens were removed, heated in 10% KOH solution and descaled, particular care 
being taken with the males not to dislodge the hairs of the subdorsal scent patches or sternal hair pencils, if 
present. As it was found that many genitalic characters would be obscured and/or distorted by the process 
of mounting, they were scored from unmounted preparations and routinely checked by dissection of a 
second example of each sex (except for Omorphina for which there was insufficient material). This was not 
done for the other characters. However, in any instance where there was doubt or ambiguity as to the 
correct coding of character states, additional specimens were examined. The numbers of each sex of each 
species studied are given in Appendix 1 . 

The method varied slightly for the six genera mentioned above, for which only very limited material (2-6 
specimens in total) was available. For five genera (not Loboplusia), the head and prothorax were not 
removed. Only one tegula, patagium and set of legs were detached for examination. The wing venation and 
structure characters were scored by wetting the wing in situ with ethyl acetate. The female frenulum was 
examined by carefully brushing the scales away at the base of the wing underneath with a 00 sable-hair 
brush. The genitalia of one male and one female only were dissected. For Loboplusia, only one set of legs 
was removed from the paratype male for examination and the genitalic characters were scored from 
pre-existing slide preparations. These measures resulted in numerous 'missing data' scores (*) in the data 
matrix. In order to reduce their number and to speed up the computer analysis, any * character that was 
found to be constant for all the other plusiine taxa was scored appropriately for the six rare genera. These 
characters were mainly from the head, prothorax, patagia and tegulae. In addition , no females of any of the 
three species of Plusiotricha (P. livida, P. carcassoni Dufay and P. fletcheri Dufay) are known and thus 
characters 14, 176, 177 and 179-216 were scored as *. Character 178, scored from females in all other 
genera, was scored from the male in Plusiotricha. Several characters also exhibited dependency upon the 
possession of certain states for other characters. These non-comparable characters were also scored as *. 
The data matrix is presented in Appendix 3. 

Scanning electron micrographs of proboscis structures were obtained using samples heated in 10% KOH 
solution, air-dried, affixed to stubs using an epoxy resin, coated with gold for 90s using a sputter coater 
(Polaron E5000), and then observed using a Cambridge S180 scanning electron microscope, with an 
accelerating voltage of 10 kV. 

Characters 

Where possible, the states of a character were arranged into an ordered transformation series. 
The states of such a series are coded (plesiomorphic), 1,2. . . n (most apomorphic). For a 
number of characters, there was uncertainty as to the most appropriate ordering of the states. 
These characters are analysed as unordered (using the UNORDERED option of the PAUP 
program; see also Analytical procedure), wherein no a priori assumptions of the order of state 
transformations is made. Unordered characters are indicated below by an asterisk and the 
coding of their states 0, 1,2 . . . n carries no connotation of polarity or transformation sequence. 



98 IAN J. KITCHING 

In several of the descriptions, the symbol ± is used to signify 'more or less' or 'approximately'. 
Scale lines on the figures are all 1 mm unless otherwise stated. 

Wing pattern 

1 Raised scales in the forewing cell absent (0)/ 'present (1). 

2 Orbicular, suborbicular and reniform stigmata outlined in black, the former two partially or totally 
fused (Figs 32, 33) (\)lstigmata not so formed (0). 

3 Lower half of suborbicular stigma, with or without a basal, exterior spot, modified to form a Y-mark 
(e.g. Figs 39, 43, 87) (\)lnot (0). 

4 *Forewing colour internal to the terminal line not pink (O)lvarying from a few scattered pink scales to a 

distinct pink line (l)/a pink line tending to be concentrated at the apex of vein M 3 (Fig. 53) (2)/a 
conspicuous pink spot at the apex of vein M 3 (Fig. 54) (2>)la distinct white line (4). 

5 Single black scales scattered over the wing surface (Figs 62, 71) {\)lnot (0). 

6 Scale tooth on the forewing tornus well developed (Figs 46, 86) (\)l weakly developed or absent (Figs 90, 
92) (0). 

7 Basal two-thirds of hindwing yellow (\)lgrey or brown (0). 

8 Base of hindwing upperside with broad scales (O)lhindwing base hyaline, scales setaceous (Figs 98, 99) 

(1). 

9 Forewing with large, discrete patches of green-gold metallic scales (Figs 47, 84, 85, 102) (\)lnot (0). 

10 Forewing with metallic sheen over parts, not arranged into discrete patches (\)lforewing without 
metallic sheen (0). 

11 Forewing with a conspicuous black spot infringe at apex of M 3 (Figs 38, 39, 51) (l)/not (0). 

12 Reniform stigma at least partially outlined in silver (Figs 58, 79, 92) (\)lnot (0). 

13 Forewing cell underside with hair-like scales (\)lwithout such scales (0). 

Wing structure 

14 * Structure of female frenulum: 

(0): 1 < (2 = 3); 1 finer and just shorter than 2 and 3 (Fig. 124); 

(1): 1<(2 = 3); 1 is finer and half the length of 2 and 3 (Fig. 125); 

(2): 1 < (2 = 3); 1 is very fine and almost lost (Fig. 126); 

(3): 7<2<3(Fig. 120); 

(4): 2 < 3; 1 lost (Fig. 121); 

(5): 3 only; 1 and2lost (Fig. 122); 

(6): 1=2 = 3 = 4; 4 setae present (Fig. 123). 
A single specimen of Pseudeva was observed with four setae on the right wing only, while a single 
Omorphina was observed with five setae on each wing. All other examples of these genera examined 
showed state 0. 

15 Hindwing vein M 2 as strong as M 3 (Fig. 131) (Q)lhalfthe width ofM 3 (Fig. 132) (l)/vestigial (no trachea 
visible internally) (Fig. 133) (2)1 lost completely (Fig. 134) (3). 

16 Apex of forewing vein R4 produced as a point beyond the apex of R 5 and M ) (Fig. 129) (I)/ not (Fig. 130) 
(0). 

17 Forewing vein R 2 + 3 + 4 (upper edge of the areole) approximately parallel to R { (Fig. 135) (0)/markedly 
converging on R } then curving away distally (Fig. 136) (1). 

18 Hindwing vein M 2 arises near M 3 such that M 2 - M 3 s£ M 3 - CuAj (Fig. 134) (0)/M 2 arises less than 
halfway between M 3 and the bend in m t - m 3 , M 2 - M 3 » M 3 - CuAj (Fig. 137) (\)IM 2 arises half 
way between M 3 and the bend in mi - m 3 , M 2 - M 3 » M 3 - CuA, (Fig. 138) (2)1 M 2 arises at the bend 
in m, - m 3 , M 2 - M 3 >» Mj-CuAj (Fig. 139) (3). 

19 Hindwing vein M 2 apex produced as a blunt point, giving five concavities between R s and CuA 2 (Fig. 
128) (0)/M 2 apex not produced, giving four concavities (Fig. 127) (1). 

20 * Scale bases on wings randomly distributed (Fig. 140) (0)/arranged in irregular transverse rows between 

the veins (Fig. 141) (l)larranged in highly regular transverse rows, one scale wide (Fig. 142) (2). This 
character is scored by observation of the scale base arrangement between forewing veins M 3 and M 4 . 

21 Forewing vein CuP absent (0)1 a distal short length of vein CuP present (Fig. 143) (1). 

Head 

22 Scales anterior to the bases of the antennae unmodified (0)/ 'modified into lashes, not exceeding the eye 
diameter (\)llashes exceeding the eye diameter (2). 



SYSTEMATICS OF THE PLUSIINAE 99 

23 Eyes large and bulbous (Figs 95, 152) (1)1 not (Figs 88, 148) (0). 

24 Prominent rim round eye absent (Fig. 148) (0)/present (Fig. 151) (1). 

25 Lower frons scaled only at the edge (I)/ 'narrow, central band offrons, at most, unsealed (0). 

26 In lateral view, frons distinctly exceeding the eye (Figs 147, 148) (0)/frons barely exceeding the eye, if at 
all (Figs 149, 150) (1). 

27 *Pilifers elongate (Fig. 144) (0)/ 'triangular (Fig. 145) (\)lvery broadly triangular, not exceeding the 

labrum (Fig. 146) (2). 

28 Ventral to antennal sockets, frons slightly concave, so giving the appearance, in front view, of a 
dorsally delimited frons (Fig. 147) (Q)/no such concavity, frons appearing undelimited ventral to the 
antennal sockets, sockets contiguous with it (Fig. 149) (1). 

29 Postocciputflat or slightly concave (Fig. 149) (0)/ distinctly convex and bulged (Fig. 147) (1). 

30 In the male, frons approximately square, as long as wide (Fig. 151) (0)/ 'distinctly narrower than long 
(Fig. 152) (1). 

Proboscis 

31 * Proboscis distally armed with no styloconic sensilla (0)/few (Fig. 108) (\)lmany (Fig. 109) (2). 

32 * Proboscis long and thin (0)/ 'shorter and stouter {1)1 reduced and non-functional (2). 

33 Styloconic sensilla with segment 1 cylindrical (Figs 110, 111) (0)/withfive longitudinal ridges, apically 
produced to give a stellate appearance when viewed end-on (Figs 112, 113) (1)1 with the three distal 
ridges produced as 'wings' (Figs 114, 115, 116) (2). 

34 * Setae on outer surface of proboscis restricted to approximately the most basal half turn (0)/ 'continuing 

almost to the tip and the section with dense styloconic sensilla (1)/ 'continuing to the apex (2). 

35 * Strengthening bars of proboscis in the most apical part semi-circular (Fig. 108) (0)/broken up into 

small, individual nodules (Fig. 117) (l)/as ( I ) but nodules only at the extreme apex (Figs. 109, 1 18) (2). 
A nodular apex (state 1) is also found in many butterflies (Sellier, 1975) and probably represents the 
plesiomorphic condition. 

36 Basal microspines on proboscis irregularly and sparsely distributed (Warranged in regular, dense, 
circumferential rows (Fig. 1 19) (1). 

Labial palps 

37 Ventral edge of segments 1 and 2 fringed with hairs (0)/ 'fringed with broad scales giving a smooth edge 
(1). 

38 Lateral scales of segment 2 adpressed (0)/ 'semi-erect (1). 

39 Ventral fringe of segment 2 (and usually 3) formed into a neat ridge, giving a blade-like appearance 
(l)lscales loose, edge not blade-like (0). 

40 Segment 1 more than two-thirds length segment 2 (Fig. 155) (0)/segment 1 approximately equal in 
length to segment 2 or slightly longer (Fig. 153) (1). 

41 Segment 3 approximately as long as segment 2 (Fig. 156) (l)lsegment 3 only two-thirds as long as 
segment 2 or less (Fig. 155) (0). 

42 Apex of segment 3 blunt (Fig. 155) (Q)l acutely pointed (Fig. 154) (1). 

43 Vom Rath's organ conspicuously papillate basally (Fig. 158) (\)lnot (Fig. 159) (0). All VROs are 
papillate in that they have individual internal invaginations, presumably housing sensilla. However, 
regular papulations, clearly visible under x 10 magnification, are termed conspicuous. 

44 Segment 3 much longer than wide (Fig. 155) (0)/subspherical (Fig. 153) (1). 

Antennae 

45 Ventral hairs on male antenna short, less than width of the segments (Fig. 162) (0)/long and apically 
very flexuous , much longer than segments are wide (Fig. 161) (1). 

46 *Male antennae simple (Figs 43, 162) (0)i 'asymmetrically bipectinate (Figs 100, 161) (\)l symmetrically 

bipectinate (Figs 92, 160, 163) (2). 

47 * Pectinations of male antenna apically with a single small seta (Fig. 161) (0)/ 'with a single large, stout seta 

(Fig. 160) (\)lwith two apical setae (Fig. 163) (2). 

48 Longest pectinations on male antennae arising from base of the segments (Fig. 163) (0)/arising distally 
on the segments (Figs 160, 161) (1). 

External structure of thorax -scaling, patagia, tegulae 

49 Large, bilobed, truncate crest present on Til (Fig. 39) (l)labsent (Fig. 104) (0). 



100 IAN J. KITCHING 

50 Tegulae adpressed to Til (Fig. 72) (0)/tegulae held almost vertically away from Til (Fig. 62) (1). 

51 Dorsal arm oftegula bluntly pointed or rounded (Fig. 166) (0)/truncate (Fig. 164) (1). 

52 * Ventral arm oftegula bluntly pointed (Fig. 171) {0)/truncate (Fig. 172) (\)l spatulate (Fig. 173) (2). 

53 Ventral arm oftegula with a colourless, microspinose extension (Fig. 169) (\)lnot (Figs 168, 170) (0). 

54 Microspines on ventral tegula arm restricted to apex and outer edge (Figs 168, 169) (0)/microspines 
extending along inner edge towards main body (Fig. 170) (1). 

55 Ventral arm oftegula produced on a lateral extension of the main body, outer edge of dorsal arm almost 
evenly continuous with the dorsal surface of the ventral arm (Figs 165, 167) (\)lventral arm not on a 
lateral extension, outer edge of dorsal arm curves back dorsally to form a distinct ridge running roughly 
parallel to the outer edge of the ventral arm (Figs 164, 166) (0). 

56 Dorsal tegula arm with a broad marginal flange (Fig. 174) (\)lflange very narrow (less than I scale base 
wide) or absent (Fig. 175) (0). 

57 Attachment point of patagium drawn out (Fig. 176) (l)/not (Fig. 177) (0). 

Legs 

58 Foretibia with a dorsal white spot (Fig. 87) (\)lwithout such a spot (Fig. 84) (0). 

59 Hind tibia and tarsi transversely banded grey, black or brown and white dorsally (Fig. 44) (0)/ 'dorsally 
unicolorous white, pale grey or pale orange (Figs 34, 87) (1). 

60 *Basitarsal comb on male hindleg absent (Fig. 178) (0)/present as a single row of short setae 

(approximately equal to the width of the basitarsus or to the length of the SV spines) (Fig. 179) (l)/a 
single row of long, fine setae (approximately 2x the width of the basitarsus or the length of the SV 
spines) (Fig. 180) (2)las (2) but setae in multiple rows (Fig. 181) (3). 

61 Inner row of SV spines on the fore-tarsi present in male on all segments along the entire length of each 
(0)/inner row ± suppressed over basitarsus (apical and subapical spines may remain) (Fig. 182) 
(l)/inner row ± suppressed on all segments, outer row suppressed on tarsi 2-5 (Fig. 183) (2)lall SV 
rows suppressed, V row of spines on basitarsus short and stout (Fig. 184) (3). 

62 Spines of hindbasitarsal SV and V rows equal along segment in bothsexes (Fig. 185) (0)/spines on basal 
half of segment smaller and finer (Fig. 186) (\)lspines suppressed on basal half of segment (Fig. 187) 
(2). For the purposes of coding character 62, the male hindbasitarsal comb of certain plusiines has 
been ignored. 

63 *Fore- and midtibiae unspined (0)/with 1-2 apical spines on outer surface (occasionally I on foretibial 

inner surface) (Fig. 195) (l)/fore- and midtibiae extensively spined on both surfaces (Fig. 196) (2). 

64 * Hindtibiae unspined (0)/spined on outer surface distal to median spur insertions (Fig. 188) (\)lspined 

along entire outer surface (Fig. 189) (2)lspined between the median and apical spur insertions 
subventrally on inner surf ace (Fig. 190) (3). 

65 * Tarsi with few small, SD and D setae (Fig. 194) (0)/with a small number of additional large spines, of 

the same form as the SV and V spines (Fig. 193) {\)lwith a few SD and D setae, finer than the SV and V 
rows (Fig. 192) (2)lwith numerous long, fine SD and D setae (Fig. 191) (3). 

66 Hindleg (measured as femur base to apex of pretarsus) subequal to midleg or longer (0)/hindleg shorter 
than midleg by approximately half the length of the mid-femur (1). 

67 Claw with a ventral lobe (Figs 201, 202, 203) (\)lclaw evenly curved ventrally (Fig. 204) (0). 

68 Hindleg inner apical spurs less than half the length of the basitarsus (Figs 199, 200) (0)/ 'approximately 
three-quarters length of the basitarsus, occasionally subequal to it (Fig. 198) (\)lspurs as long as 
basitarsus or longer (Fig. 197) (2). 

69 Foretibia with a dorso-apical claw (Figs 205, 206, 207) (\)lclaw absent (0). 

Internal structure of prothorax 

70 Dorsal arms offurca vertical, not apically produced (Fig. 209) (0)/dorsal arms produced and curved 
posteriorly (Fig. 210) (1). 

71 Dorsal edge of epimeron with a dorso-ventral ridge running from near the dorsal f ureal arms to the 
apex of the SV f ureal arms (Figs 212, 213) (\)lwithout such a structure (Fig. 211) (0). 

Male abdomen A 1-7 

72 Abdomen cylindrical or broadly tapered (0)/ 'abdomen distinctly narrowly tapered, A 7 half the width of 
A2(\). 

73 * Internal flange on anterior edge of T2 curved and of uniform diameter (Figs 216, 219) (0)/with two SD 

lobes (Fig. 220) (\)llobes almost fused but with a central emargination (Fig. 221) (2)/flange semicircu- 
lar, ventral edge straight (Figs 217, 218) (3). The flange in Cucullia is broad but uniformly so and with a 



SYSTEMATICS OF THE PLUSIINAE 101 

distinctly concave ventral edge thus distinguishing it from state 3 in which the ventral edge is straight. 
The flange in Lophoptera is reduced but still has a straight ventral edge, as well as the characteristic 
triangular dorsal inflection. The latter structure also serves to distinguish state 3 from extreme forms 
of states and 2. 

74 Central membranous area of St2 flanked by grooves arising from anterior apophyses or by flat cuticle 
(Fig. 222) (0)/flanked by two internal longitudinal flanges arising from the bases of the anterior 
apophyses, and which are directed obliquely towards the midline (Fig. 223) (1). 

75 * All components of triflne basal abdominal hair pencils absent (Q)/some components present (\)lall 

components present (2). Of the five main components of the trifine hair pencils (Birch, 1972a), 
Magusa possesses only the lever and the pockets, and these are reduced and non-functional. Because 
this condition could be the result of partial gain or loss, the character is treated as unordered. 

76 St7 broad, ± entire (Fig. 208) (0)/St7 with two postero-lateral sclerotizations, giving a 'fish-tail' 
appearance (Figs 214, 215) (1). 

77 Dorsal-lateral hair tufts on A2-7 absent (0)/present on A5-7 (Figs 46, 57) (1)/ 'additional 'secretory' 
areas present on A4, occasionally on A2-3 (2)1 distinct additional tufts present on A4 (3) . The secretory 
areas on A2-4 are very friable and are easily lost during preparation. Thus no attempt was made to 
subdivide state 2, especially as it would not be possible to reliably score Eutheiaplusia and 
Plusiopalpa. Loboplusia has a unique distribution of what may be the scale bases of scent tufts on the 
pleura of A2-4 but this is uncertain from the descaled preparation. The genus was thus coded as *. 

78 * Scent hairs on A7 exposed (Fig. 226) (0)/ 'enclosed in an invaginated groove (Fig. 227) (l)lenclosed in an 

overfold of pleuron 7 (Fig. 228) (2)lenclosed in a membranous fold (Fig. 229) (3). These three states 
are probably independent derivations. 

79 * Dorso-lateral hair tufts on A5-6 with indistinct edges (0)/ 'tufts on A5-6 with distinct, sharp antero- 

medial edges (\)las (1) but A5 only (2)las (1) but A6 only (3). 

Male SIS and the ventral hair pencils 

80 *St8 with a thin anterior bar and two broad lateral bars extending posteriorly (Fig. 235) (0)/lateral bars of 

St8thin (Fig. 243) (\)las (1) but anterior bar narrowed medially, often with latero-ventral, posteriorly- 
directed sclerotized flaps (Fig. 254) (2)llateral arm of St8 produced at an acute angle to the anterior arm 
(Figs 236-240) (3). This character probably comprises two independently derived forms of St8 and 
hair pencil: the plusiine type, states 1 and 2; and the stictopterine type, state 3. The plusiine type is 
probably derived directly from state but this is much less certain for state 3. Subsequently, the two 
types of hair pencil and associated St8s will be treated as mutually exclusive. Characters applicable to 
one type will be coded in taxa showing the other type as * . The same coding will be used for those taxa 
showing state for 80. Several taxa exhibit partial reductions of the hair pencils and St8. Cucullia and 
Abrostola have much reduced lateral arms but still can be interpreted as 80(3) and 80(1) respectively. 
In the following hair pencil characters, 81-82 refer to the stictopterine type, while 83-95 refer to the 
plusiine type. 

81 * Hair pencils arising from a diamond-shaped pocket posterior to the anterior arm ofSt8 (Figs 238, 239) 

(0)/from two shallow ventral pockets (Fig. 240) (\)lfrom two deep lateral pockets (Fig. 237) (2)lpencils 
reduced to a ventral row of scales arising from a very thin sclerotized bar running just posterior to and 
parallel with the anterior arm of St8 (Fig. 236) (3). These four forms of hair pencil appear to be 
mutually exclusive. 

82 Angle formed by anterior and lateral arms of St8 without an exterior lobe (Fig. 236) (0)/with a small 
exterior lobe (Fig. 238) (\)lwith a large, dorsally recurved bar (Fig. 239) (2). 

83 * Hair pencils poorly developed, merely a tuft of sparse scales arising from a shallow depression (Fig. 

241) (0)/hair pencil pockets slightly invaginated but still not discrete (Fig. 245) (\)lpockets well 
developed, partially fused along their length (Fig. 243) (l)lpockets well developed, separate along their 
entire length from St8 to their apices (Fig. 244) {i)lpockets well developed, totally fused into a single 
structure but the two individual pencils still discrete (Fig. 265) (4)/single pocket present, elements fused 
(Figs 242, 248) (5). It is uncertain whether the undeveloped hair pencils of Abrostola and of 
Polychrysia and its relatives are plesiomorphically undeveloped or the result of subsequent reduc- 
tion. All these genera were coded 0. 

84 Hair pencil pockets with a sclerotized bar along the inner ventral edge (Figs 254, 258) (1)/ 'absent (Figs 
256, 263) (0). 

85 Hair pencil insertions in one main apical field (Figs 255, 262) (0)/ hair pencil insertions differentiated 
with distinct, discrete subdorsal and lateral fields (Figs 258-260, 269-271) (1). 

86 Hair pencil pockets with small subventral basal accessory pouches (Figs 260, 263) (1)/ 'absent (Fig. 257) 
(0). 



102 IAN J. KITCHING 

87 Hair pencils composed of scales of one length (Fig. 257) (0)/ 'outer scales markedly shorter than inner 
scales (Figs 263, 264, 267, 268) (1). 

88 Hair pencils with an inner tuft that is much longer than the remaining inner scales of the pencil (Figs 
260, 272, 273) (\)lall inner scales of approximately equal length (Fig. 266) (0). 

89 Lateral arms of St8 bearing secondary hair brushes (Figs 242, 261, 267) (l)/not (0). 

90 Lateral arms of St8 with an anterior flange where they curve dorsally over the hair pencils (Figs 260, 
271) (1)/ 'without such a flange (Fig. 264) (0). 

91 Lateral arms of St8 undifferentiated, running to posterior margin of segment 8 as uniform linear bars 
dorsal and lateral to the hair pencils (Fig. 243) (0)/lateral arms twisted dorsally over the hair pencils 
(Fig. 257) (\)las (1) but also with a deep basal concavity in outline (Fig. 262) (2). 

92 * Lateral arms of St8 roughly linear (Fig. 244) (0)/with a broad triangular exterior sub-basal lobe (Fig. 

256) (\)llobe modified into a blunt projection (Fig. 258) (2)/lobe attenuate and recurved dorsally (Fig. 
265) (3). 

93 * Dorsal arms ofSt8 run parallel and do not meet apically (Fig. 245) (0)/ 'dorsal arms converge to a point 

(Fig. 254) (l)ldorsal arms run approximately parallel, fusing into a rounded apex (Fig. 263) (2)/ 'dorsal 
arms run transversely, approximately at right angles to the long axis of the body (Figs 258, 269) (3). The 
dorsal arms are missing in Eutheiaplusia, the genus being coded * for those characters (93-94) 
dependent upon their presence. 

94 * Ventrally, dorsal arms ofSt8 unsealed or with unmodified scaling (0)/with broad, spatulate, erect scales 

(Fig. 242) (\)lwith broad, flat, 'greasy-looking scales, oppressed to the surface and adhering into sheets 
(2)/with scales as in (2) but those apically on the dorsal arms formed into a semi-circular flat plate 
projecting distally (Figs 258, 269) (3). 

95 Membrane between lateral arms of St8 ventrally bare (0)/covered in long, fine, hair pencil-like scales 
(1). These hairs are very easily brushed off, which may be what has happened in the genera 
Antoculeora, Macdunnoughia and Allagrapha, in which the hairs are very sparse but nevertheless 
recognizable. 

Male genitalia - uncus, tegumen & anal tube 

96 Uncus a simple, ± cylindrical tube (Fig. 284) (0)/uncus flattened ventrally so it is D-shaped in 
cross-section (Fig. 286) (1)/ 'uncus reduced to a subspherical structure (Figs 276, 277) (2). This character 
is treated as unordered. The three types are treated as independent for appropriate subsequent uncus 
characters, as was done for hair pencils. It should be noted that there are numerous modifications to 
state 0, some of which are coded below as separate characters. 

97 Uncus basally curved (Fig. 282) (0)/straight (Fig. 283) (1). State can show considerable variation 
(Figs 281, 282, 284, 285). 

98 Uncus of even width or gradually tapering distally (Fig. 284) (0)/uncus distinctly distally dilated 
dorsally (Fig. 293) (1). 

99 In lateral view, dorsal margin of tegumen ± continuous with dorsal margin of uncus (Fig. 282) (0)/a 
distinct notch present between tegumen and uncus (Fig. 285) (l)/apex of tegumen produced, apical 
angle about 90 degrees (Fig. 287) (2). In state 1, the notch is formed by the downcurvature of the apex 
of the tegumen and the increased basal curvature of the uncus. In state 2, the tegumen curvature is 
accentuated to a sharp angle. Abrostola is superficially state 2 but the tegumen is apically rounded 
(see Fig. 274). State may be flat or have a uniform convex curve (Figs 281, 284). Several plusiines 
(e.g. Trichoplusia, Dactyloplusia and Palaeographa) have a reduced state 1 that superficially looks 
like state 0. 

100 Base of uncus convex (Fig. 285) (0)/concave (Figs 274, 279) (1). 

101 In lateral view, dorsal edge of tegumen, distal to pleurite, evenly curved or straight, with at most a short, 
flat apex (Fig. 281) (0)/with a distinct angle, just over halfway along; when origin of uncus is horizontal, 
the distal section of tegumen dips towards uncus base (Fig. 285) (1). Many other genera have flat distal 
sections to the tegumen (e.g. Diachrysia, Chrysodeixis, Palaeographa) but these are all horizontal 
when the uncus base is aligned as described above. 

102 Uncus medially cylindrical {0)/medially slightly to greatly dorso-ventrally flattened and laterally 
expanded (Figs 275, 278) (1). 

103 In dorsal view, uncus with a distinct median suture visible, especially basally (Figs 275, 278) {\)l absent 
(Figs 288, 290) (0). 

104 In dorsal view, outer edges of tegumen diverge at about 90 degrees or less (Figs 288, 290, 296) (0)/outer 
edges diverge at almost 180 degrees (Figs 295, 297) (1). 

105 Dorsal edge of anal tube arising from base of uncus or extreme apex of tegumen (Fig. 282) (0)/ 'anal tube 



SYSTEMATICS OF THE PLUSIINAE 103 

arising from bases of latero-apical sclerites of the tegumen (LASTs) to about halfway along the LASTs, 
free of uncus base (Fig. 284) (1). 

106 LASTs free apically of ventral edge of tegumen, projecting to articulate with the uncus base (Fig. 275) 
(0)/ LASTs fused to underside of tegumen, ± fused dorsally around uncus base (Figs 290, 296) (1). 
Also, in Paectes, the LASTs extend ventrally under and fuse under the anal tube as a sharply pointed 
structure, a true gnathos. In their reduced form, the plesiomorphic state consists of a pair of lateral 
bars extending from the ventral margin of the tegumen to the uncus base. In the plusias, the 
membranous portion becomes sclerotized, either completely or leaving only a small, circular 
membranous area basal to the uncus medially. In some weakly sclerotized genera (especially 
Polychrysia and its relatives), there may be subsequent desclerotization but this is general and the 
plesiomorphic condition is not displayed. Plusiopalpa dichora has extensive membranous areas 
centrally but overall is like the other plusias. LASTs are absent in Anuga. 

107 Dorsal surface of anal tube membranous (0)/with two subdorsal bars arising from the LASTs (1). 
These bars are very weak in Abrostola. In Stictoptera, Lophoplusia, Lophoptera and Chrysanympha, 
the connection with the LASTs is tenuous, while the bars in the latter two genera are fused into a 
single dorsal bar. The bars in Mouralia bear subdorsal, wing-like projections. 

Male genitalia - vinculum & pleurite 

108 Vinculum arms with a small, distinct, ventral tooth (Fig. 303) (Y)lwithout a tooth (Fig. 308) (0). 

109 In lateral view, vinculum apically straight or with only a single slight curve (Fig. 299) (0)/vinculum with 
two apical curves, dorsally then ventrally (Fig. 319) (1). 

110 Arms of vinculum ± straight in ventral view, vinculum V-shaped (Fig. 303) (0)/arms bent about 
halfway along, vinculum Y-shaped (Fig. 301) (1). State 1 shows a distinct Y-shape, with the anterior 
parts of the vinculum arms running ± parallel. In Polychrysia and several other genera, the vinculum 
is slightly Y-shaped but the distal parts still markedly converge. 

111 Vinculum apically produced as a long, thin spine (Figs 300, 308) {1)1 vinculum not so produced (0). 
Both Pseudoplusia and Chrysodeixis have vincula produced to a sharp point. In Stigmoplusia, the 
vinculum apex is rounded. 

112 * Dorsally andsubapically, dorsal flanges of the vinculum arms with even edges (Fig. 303) (0)/edges with 

two small emarginations towards the exterior (Fig. 298) (\)l emarginations produced into two sharp 
points (Figs 299, 300) (2)1 emarginations produced into two blunt posteriorly-directed hooks (Figs 304, 
305) (3). 

1 13 Dorsal and ventral flanges of vinculum arms greatly expanded and flattened dorso-ventrally forming a 
broad flat cone, unfused medially, sides of vinculum subparallel (X)lflanges of vinculum arms not so 
enlarged (0). The apomorphic state for this character is a complex one that has to be qualified. The 
subparallel sides are also found in those genera exhibiting state 1 of 110 but they do not have 
expanded and flattened flanges. In Lophoptera (Fig. 309), the flanges are expanded and flattened, 
but they are medially fused to form a complete cone and the edges are distinctly convergent. 

114 Dorsal and ventral flanges of vinculum arms separate or, at most, ventral flanges fused (0)/ 'dorsal and 
ventral flanges fused across to give the appearance of a flattened cone (Figs 302, 309, 310) (1). In 
Eosphoropteryx, the ventral flanges only are fused across. In Diloba, the dorsal flanges may be fused 
but the general desclerotization makes this difficult to discern. The vinculum of Stictoptera is highly 
derivative and although what is left of the flanges appear to be fused, a complete conical structure is 
not found and the condition is not strictly comparable. Stictoptera was scored *. The fusion in Paectes 
is drawn out but a conical form is still discernible. 

115 Vinculum arms divergent to the base of the tegumen (Fig. 303) (0)/arms divergent and then run parallel 
towards, or converge towards, the tegumen (Fig. 310) (1). 

116 Pleurite dorsally simply concave, ventrally entire or with a desclerotized nick (Figs 317, 318) 
(0)/pleurite dorsally strongly concave basally and then with a sharp convex bend caused by a distinct 
sharp ventral basal notch (Fig. 312) (1). In Chalcopasta and Oncocnemis, there is a ventral basal 
desclerotization but this is not manifested dorsally by a sharp bend. In Omorphina, there is a 
dorso-basal bump but no ventral notch. 

117 Pleurite without a lateral flange, lateral margin somewhat recurved towards the midline (Fig. 306) 
{0)/pleurite with a lateral, often broad, flange, lateral margin flat (Figs 311 , 319) (1). In Anagrapha and 
Palaeographa, the lateral flange is somewhat reduced in width but the margin is nevertheless flat. 
Pseudoplusia and Chrysodeixis have very thin pleurites with no lateral flange (or one that is very 
narrow). The pleurite of Anuga is reduced to a small triangular lobe. In Paectes, a pleurite-like flange 
is fused onto the antero-dorsal margin of the tegumen (see character 123), while in Diloba, a thin 
sclerite, presumably the pleurite, occupies a similar fused position. In the stictopterines, the pleurite 



104 IAN J. KITCHING 

is either reduced to a small lobe (Stictoptera) or lost (Lophoptera) , or fused with the tegumen and the 
vinculum connection lost. 

118 Apex ofpleurite recurved dorsally over the tegumen to articulate on the inner edge in a pocket (Fig. 314) 
(0)/ 'pleurite apically lying dorsally on the tegumen or slightly recurved over the inner edge, but not 
articulating in a pocket (Fig. 313) (1)/ 'tegumen rotated almost vertically, pleurite apex only in contact 
with the tegumen, otherwise lying parallel with the ventral apical vinculum bar (VAVB) (Fig. 316) (2). 
In state 2, the pleurite and the VAVB appear as linear continuations of the vinculum apex. 

119 Pleurite and VAVB fused basally (Fig. 312) (0)1 fused to about half the length of the pleurite (Figs 306, 
307) (1). 

120 Pleurite broad, curved in lateral view and separate from the antero-dorsal margin of the tegumen (Fig. 
312) (O)Zpleurite very narrow, very closely oppressed to (but not fused with) the antero-dorsal margin of 
the tegumen (Fig. 289) (1). 

121 *VAVB a long linear bar projecting from the apex of the vinculum to articulate ventro-apically on the 

tegumen, apex of vinculum and base of tegumen distinctly separate (Fig. 312) (0)/VA VB very short or 
absent, tegumen and vinculum closely approximated (Fig. 311) (l)/VA VB absent or reduced to a blunt 
lobe, tegumen and vinculum separated (Figs 318, 321) (2)/VAVB absent, tegumen produced basally 
below pleurite to approach vinculum apex (Figs 315, 317) (3). In state 1, if the VAVB is present, then 
it is bent, thus allowing the vinculum and tegumen to come into close proximity; in state 0, the VAVB 
is straight. 

122 VAVB produced ventro-posteriorly into a large, articulated socius (Figs 291, 292) {\)lnot (0). In 
Paectes, the socii are hairy, unsclerotized, blunt lobes; in Amiga, they are long, sclerotized, curved 
and apically hooked, and are remarkably convergent in appearance upon the typical plusia uncus. 

123 Vinculum and tegumen separate or linked by the VAVB (0)/ 'vinculum and tegumen broadly joined, 
pleurite totally fused along the antero-dorsal edge of the tegumen (Fig. 320) (1). This character could 
also be represented as yet another alternative state of character 121. However, given that the 
euteliines and stictopterines have been tentatively coded for character 121 (as opposed to * for 
Diloba), characters 121 and 123 have been retained as separate. 

Juxta & anellus 

124 Anellar membrane present between ventral margin of aedeagus and apex of juxta (Fig. 323) (0)/apex of 
juxta ± fused to apex of aedeagus ventrally (Fig. 322) (1). 

125 Apex of juxta uniformly thin (0)/ 'apex with a transverse apical thickening and a median longitudinal bar 
(Figs 324, 325) (1). The longitudinal bar in Abrostola is weak, while in Mouralia, it is apparently 
double. 

126 Juxta free from valve latero-basally (0)/ 'juxta fused to inner edge of sacculus near the base of the clavus 
by a short thin bar (Figs 324, 325) (1). 

127 Apex of juxta entire or if decurved, no notch or apical process present (Figs 325, 329) (0)/ 'apex folded 
ventrally to form a ± parallel-sided notch that may be partly sclerotized dorsally to form an apical 
process (Figs 326, 327) (\)lprocess well sclerotized distally so as to appear subapical on the juxta (Figs 
335, 336, 340) (2)1 process further modified into a sharp upcurving spine (Figs 337, 338, 339) (3). 
Thysanoplusia, Zonoplusia and Ctenoplusia have juxtas in which the apex comes to a point without 
curving ventrally. Similarly, in Trichoplusia, Acanthoplusia, Agrapha and Anadevidia, the juxta is 
apically deeply concave but not decurved. Also, in these six genera, the anellus arises apically from 
the juxta. In those genera showing states 1-3, the anellus arises dorsally and subapically from the 
juxta, due to the dorsal sclerotization of the apical process. 

128 Apex of juxta smooth (0)lapex of juxta scobinate (Figs 328, 329, 330, 334) (\)lscobinations distally 
modified into sclerotized thorns (Fig. 341) (2). The scobinations in Chrysodeixis are very weak, while 
in Pseudoplusia, they are restricted to the lateral, distally-produced arms of the juxta. Thus, the 
states in these two genera may not be homologous with that in Dactyloplusia and Thysanoplusia. The 
apices in Plusiopalpa and Cucullia are also heavily scobinate and even spinose (Plusiopalpa shisa), 
but in these genera the juxta merges gradually into the anellus and so it is difficult to say on which 
structure the ornamentation is located. Nevertheless, they were both coded 1. 

129 Juxta a ± quadrate sclerite, and usually well sclerotized (0)1 juxta crescent- or boomerang-shaped, 
poorly sclerotized (Fig. 333) (1). The juxta of Polychrysia and its relatives is poorly sclerotized but 
distinctly quadrate. Stictoptera has a highly autapomorphic juxta and is coded *. The juxtas of 
Brachionycha and Diloba are somewhat crescentic but are well-sclerotized. 

130 Juxta basally separate from saccular flanges (Fig. 342) (0)/juxta basally fused to basal saccular flanges 
(Fig. 343) (1). Those genera lacking saccular flanges (see character state 131:0) are coded *. 



SYSTEMATICS OF THE PLUSIINAE 105 

Valve 

131 Base ofsacculus evenly rounded (Figs 372, 373, 374) (0)/with a basal flange (Figs 351-355) (1). 

132 Saccular flanges fused across and produced medio-anteriorly into a rounded lobe between the valve 
bases (Fig. 343) (\)lflanges not so modified (0). In most genera with saccular flanges, they are either 
separate from each other or fused across to form an irregular flat area of sclerotization. In two genera 
(Panchrysia and Chrysanympha), however, the flanges are formed medially into a rounded, 
somewhat dorsally concave lobe, giving, with the valve bases, a regular anterior outline like a 
rounded M or a two-cycle sine wave. 

133 Ventral edge of valve basally with a single row of enlarged flattened blade-like setae (Figs 351, 355) 
(I)/ without such setae (0). 

134 Sacculus, distal to clavus, produced into an irregular spatulate lobe (Figs 347, 348) (\)lnot (0). This 
structure has been widely interpreted in the past as the clavus. However, its position (more distal on 
the sacculus margin than the position occupied by the clavus in other noctuids) and the presence of an 
obvious clavus in the 'correct' location in Mouralia argue that the spatulate lobe is not the clavus but a 
novel development apomorphic for the Abrostolini. 

135 Apex of valve with a distinct uniseriate corona (Fig. 349) (\)labsent (0). The corona is defined here as a 
distinct marginal series of strong basally-directed spines. In Calophasia and Chalcopasta, the corona 
is weak and somewhat submarginal. 

136 Apex of valve not differentially scaled (0)/with multiple rows of easily detached, basally-directed, setose 
scales; when removed, scale bases forming a distinct field (Figs 351-355) (1). 

137 Marginal setae on ventral edge of valve on normal pinacula (Figs 347-350) (0)/some setae arising from 
marginal prominences (Figs 368, 369, 370) (1). The marginal prominences are much larger than 
normal pinacula and frequently bear two or more setae. 

138 Valve constricted at about two-thirds of the distance base to apex, apical section spatulate (Figs 354, 
355) {l)/valve not constricted (Figs 379, 382) (0). 

139 Dorsal edge of saccular overfold with a lobe that overlies and obscures the clavus base when the valves 
are opened flat (Figs 360, 368, 369, 370) (\)lclavus base, if present, not obscured by sacculus when 
valves opened flat (Figs 348, 361) (0). 

140 Ventral edge of valve basally evenly curved (0)/ 'with a broad, blunt point about one-third from base 
(Figs 351, 352) (1). 

141 Sacculus with a broad, triangular spine basal to the origin of the harpe (if present), often further 
modified (Figs 370, 376, 378) (\)labsent (0). This saccular spine is modified into a quadrate lobe in 
Erythroplusia and into a long, serrate, lance-like projection in Antoculeora. 

142 Apex of sacculus continuous with ventral margin of valve (0)/apex produced as a sheet, costa extended 
ventrally towards it, but not fused with it (Figs 368, 369, 370) (l)lsaccular sheet and costal overfold 
fused, often modified into a curved hook (Figs 361, 372, 373, 374, 376) (2). This character is highly 
problematical given the high degree of structural diversity displayed by the genera that possess it. 
Allagrapha and Macdunnoughia exhibit the simplest form in which the apex of the sacculus is 
extended beyond the ventral margin of the valve as a broad, blade-like sheet. This is modified in 
Puriplusia into an inwardly-curving hook. In Autoplusia, a condition similar to that seen in Puriplusia 
is found, in that the apical saccular process is modified to form an inwardly-curving hook. However, 
in Autoplusia, there is a distinct fusion with the costal margin in the formation of the hook that does 
not appear to occur in Puriplusia. Sclerogenia is similar to Autoplusia but the hook is larger, apically 
blunt, and is curved ventrally rather than inward, while the ventral edge of the valve is greatly 
enlarged, giving the hook an apparently subdorsal origin on the valve. The valve of Antoculeora is 
highly autapomorphic and very difficult to homologize with the valves of the other genera. I have 
chosen to interpret the ventral valve digitations, and in particular the subapical one, as being 
homologous to the saccular hook of Autoplusia and Sclerogenia. However, given that the inner 
margins of the costa and sacculus in Antoculeora are completely fused along almost their entire 
length, a more comprehensive study is required to confirm this and the previous interpretations of 
this character. 

143 * Median edge ofsacculus basally unmodified (0)/produced into a broad, rounded, setose lobe (Fig. 356) 

(l)/lobe elongated into a distinct clavus, dorsal and ventral margins of which are basally ± level (Fig. 
357) (2)1 dorsal margin much cut back (or ventral section basally elongate) so that clavus appears to be 
located on a sclerotized peninsula (Fig. 358) (3). 

144 Clavus a distinct tubular structure, >3x as long as basally wide (Fig. 360) (0)/ 'clavus a short tubular 
structure, aboutaslongas basally wide(Fig.380)(l)/clavus reduced tonomorethanaslightsetosebumpon 
the end of the peninsula (Fig. 382) (2). Thischaracter and 145 are dependent upon states of character 143. 
Character 144iscoded* for taxashowingstatesOandl for character 143, while character 145 iscoded* in 



106 IAN J. KITCHING 

taxa showing statesO, 1 and 2 for character 143 . Character 144 is included because it is one of the 'classic' 
plusiine characters and is the one used by McDunnough (1944) to divide the plusias into three groups of 
genera. Cornutiplusia and Polychrysia both have fairly short clavi but still long enough to be coded 0. 
Similarly, Euchalcia and Lamprotes are somewhat intermediate between states 1 and 2 but given that 
some tubular structure can be detected, they were coded 1. 

145 Clavus borne on a short peninsula (Fig. 358) (O)lclavus borne on a highly elongate, somewhat flattened 
peninsula that is about as long as the tubular clavus (Fig. 359) (1). Genera possessing short or very 
reduced clavi (character 144: 1 & 2) are not strictly comparable and were coded 0, since in these 
genera the peninsula is not elongated, but is longer than the clavus by virtue of the latter's reduction. 

146 Valve overall blade-shaped, ventral edge ± straight for about two-thirds of its length, then curved up to 
the apex of the costa to form a blunt or sharp point (Fig. 381) (\)lapex of valve ± evenly rounded (0). 
The shape of the valve coded by state 1 is readily recognized but difficult to describe and delimit 
briefly. The general impression is of a knife blade ; the valve gradually widens from the base , the costa 
is slightly concave and the ventral margin possesses well-developed, sclerotized setae. In Cucullia, 
the dorsal and ventral edges of the valve are ± parallel and the genus was coded 0. The valves of 
Diachrysia chryson are less blade-like than most genera coded 1, but other species in this genus, 
notably D. nadeja, are as extreme as the autographas. Anagrapha is interpreted as state 1, due 
allowance having been made for the highly modified, excavated apex in this taxon. 

147 Valve ± oval in outline, roughly symmetrical along the midline (Fig. 350) (l)/valve not oval, 
asymmetrical along the midline (0). Euchalcia variabilis has a somewhat quadrate valve but is still 
coded 1. Most other Euchalcia species have simple, oval valves (Dufay, 1968). Note that states 1 for 
characters 146 and 147 are mutually exclusive. Taxa showing state 1 for one character are coded * for 
the other. 

148 Harpe present (\)labsent (0). If the saccular processes Autoplusia, Sclerogenia and Antoculeora are 
reinterpreted as harpes, then states 1 and 2 of character 142 are not part of the same transformation 
series. 

149 Harpe a blunt, tubular process (0)/harpe modified into a sharply pointed, basally curved hook (Figs 
354, 367, 379, 380, 386) (1). A number of genera have non-tubular harpes (e.g. Lophoptera, 
Calophasia, Macdunnoughia) that are also not of the form of a curved hook (see also character 150). 
Because they do not show the apomorphic condition of character 149, these genera will be coded 0, 
even though they do not possess the plesiomorphic condition for this character either. The same 
rationale applies to the following character also. The hooked harpes of Syngrapha, Caloplusia and 
Palaeographa are almost certainly convergent upon those of Trichoplusia, Dactyloplusia and 
Agrapha, but a wider survey of species needs to be performed before the two (or more) forms here 
coded as state 1 can be adequately distinguished. 

150 Harpe a blunt, tubular process (Q)/harpe a strongly dorsally and inwardly curving hook (Fig. 369) 
{\)lbase of harpe also with an upcurving shorter hook forming a pincer-like structure (Fig. 370) (2). 
This hooked modification of the harpe curves towards the midline when the valves are closed, 
whereas that in character 149 curves in the plane of the valve towards the valve base. 

151 Valve apically entire (0)/valve apically deeply cleft, costa and sacculus separate (Fig. 387) (1). 

152 Valve much longer than wide (0)/valve ± as long as wide, overall outline somewhat trapezoid (Fig. 375) 
(1). 

153 When closed, valves lie in a lateral position, easily opened flat (Fig. 389) (0)/when closed valves lie 
ventrally or subventrally, difficult to open flat (Fig. 388) (1). 

154 Transtillae fused dorsally above aedeagus and produced posteriorly as a broad, blunt, triangular point 
(Fig. 346) (1)/ 'transtillae separate or, if fused, not produced as a broad, blunt, triangular point (Fig. 
344) (0). The euteliines lack transtillae and are coded * for this and the next character. 

155 Transtillae a single bar, though often curved, angled or broadly triangular (Fig. 344) (0)/transtillae 
laterally bifurcate into two thin bars diverging approximately at right angles; one passes dorsally over 
the aedeagus and fuses with its opposite number, the other runs lateral to the aedeagus (Fig. 345) (1). 
Omorphina has thin, fused bars passing over the aedeagus but no lateral bars. Allagrapha has broad, 
ill-defined lateral bars that may be homologous to the thin, heavily sclerotized, well-defined bars in 
Sclerogenia and Erythroplusia, but more species need to be investigated to confirm or refute this 
possibility. 

Aedeagus 

156 Aedeagus large and thick (0)/ 'aedeagus very thin and small, relative to the rest of the genitalia (1). This is 
a 'Gestalt' character, very difficult to define objectively but easy to recognize in the apomorphic 
state. 



SYSTEMATICS OF THE PLUSIINAE 107 

157 * Cuticular simplex arising dorso-basally on aedeagus, coecum ± absent (Fig. 390) (0)/cuticular simplex 

arising distinctly sub-basally , coecum distinct and of same diameter as aedeagus (Fig. 391) (\)lcoecum 
distinct and bulbous (Fig. 392) (2). In some plusiines, the bulbous coecum is only just discernible as a 
slight increase in the diameter over the apical diameter (e.g. Antoculeora) . In those genera exhibiting 
state (e.g. Oncocnemis), the aedeagus in ventral view is of constant diameter (Fig. 393) and in 
lateral view, the ventral basal curvature is often more gradual than the dorsal basal curvature (Fig. 
390] I (e.g. Cucullia). 

158 Aedeagus generally sclerotized (Fig. 393) (0)1 sclerotization apically restricted to a ventral band that 
gradually broadens onto the coecum (Fig. 394) (\)l sclerotization restricted to a thin ventral band along 
the entire length of the aedeagus distally, broadening somewhat abruptly onto the coecum (Fig. 395) 
(2). Also, the ventral band of state 2 is extremely well defined laterally, the edges being straight and 
parallel. The rest of the aedeagus is entirely membranous. In state 1, the ventral band merges 
somewhat into the membranous area laterally. 

159 Vesica ± covered for entire length in small, basally -directed spines (l)/vesica not so ornamented (1). 

160 When inverted, vesica very strongly and distinctly serpentine (Fig. 406) (l)lvesica not folded thus when 
inverted (0). 

161 Base of ductus ejaculatoris with a small, oval, sclerotized plate (\)l without (0). 

162 Vesica with an isolated, subapical cornutus (SAC) (1)1 without (0). Paectes has two cornuti, one of 
which could be interpreted as subapical but, as the form of the vesica is short and inflated, rather than 
long and tubular, as in the plusiines, Paectes was coded 0. Caloplusia has a spinose, sclerotized plate 
in the approximate position of a SAC but was coded 0. Autoplusia bears a large, lateral cornutus 
about halfway along the vesica that could be interpreted as a SAC. This would, however, require that 
either the SAC has moved from a subapical to a median position on the vesica or that a large portion 
of the ductus ejaculatoris has become inflated and gained a large field of numerous small cornuti. It is 
more parsimonious to interpret Autoplusia as having no SAC. Similarly, Palaeographa has a 
recurved hook about halfway along the vesica that also is not homologous to a SAC. A SAC was 
reported and illustrated in Eutheiaplusia by Dufay (1970a) but was absent from the specimen 
examined in this study. The genus was coded 0. SACs were also occasionally absent in specimens of 
Zonoplusia and Polychrysia. In Rachiplusia, the SAC is double. 

163 SAC apically swollen (Figs. 408, 409) (\)lapically tapered (Fig. 403) (0). In Adeva, the SAC is short 
and therefore somewhat stubby. It is nevertheless not apically swollen. Taxa coded for character 
162 were coded * for characters 163-166. 

164 SAC arising from a circular, concave plate that has a somewhat irregular margin (Figs. 362-365) 
(\)l SAC arising from vesica without a differentiated, plate dike base (0). 

165 SAC arising apically from a lateral diverticulum (1)1 SAC arising directly from the main vesica tube (0). 
The SAC of Plusia arises at the base of a diverticulum and was coded 0. 

166 SAC tubular (Fig. 403) (0)/SAC dorsally grooved (Fig. 404) (Wedges of groove produced dorsally as 
'wings' (Fig. 405) (2). 

167 Vesica with a membranous sac invaginated from the apex of the basal RHS vesica sclerotization and 
projecting anteriorly within the vesica (Figs 397, 401) (l)lvesica without such a sac (0). This sac is the 
'bulbous lock' of Callahan (1960). In Trichoplusia, Thysanoplusia, Argyrogramma and Stigmoplu- 
sia, the sac is small and ± fused to the vesica sclerotization internally. In Plusiotricha, the sac is very 
long, reaching internally almost to the apex of the coecum. 

168 Base of vesica membranous and smooth (0)/ 'internally granular, often forming a distinct annulus (1). 

169 Basal vesica sclerotization apically confluent with the membranous part of the vesica (Fig. 397) 
(0)/basal sclerotization produced ventrally as a blunt, apical point (Figs 398, 399) (\)lapical point 
elaborated into a large, blunt, recurved hook (Fig. 407) (2). The recurved hook in Abrostola is 
subventral on the LHS, rather than ventral but is interpreted as a further development of the ventral 
blunt point found in Mouralia. 

170 Vesica with a large, RHS, membranous diverticulum distal to the basal sclerotization bearing afield of 
cornuti on its outer-distal surface (\)l without such a diverticulum (0). In Mouralia, the field of cornuti 
is reduced to a small, sclerotized plate with several short cornuti. 

171 Vesica sub-basally with a dorsal, isolated patch of broad, curved, short, pointed cornuti (Fig. 396) 
(\)lwithout (0). In Lamprotes, the cornuti are connected to a LHS, lateral, sclerotized bar. Similar 
features are found in other genera (e.g. Antoculeora, Sclerogenia) , but the location and apparent 
origin do not appear to be the same as for the cornuti here coded by state 1. 

172 Vesica with two short, straight, pointed cornuti dorsally distal to the apex of the basal sclerotization 
(Figs 397, 401) (l)lwithout (0). 

173 Basal sclerotization of vesica on RHS uniform (0)/modified and reduced to a lateral, longitudinal band, 



108 IAN J. KITCHING 

differentially sclerotized dorsally and ventrally (Fig. 400) (1). Plesiomorphically, the base of the 
aedeagus appears to have been ± uniformly and heavily sclerotized, forming a basal, permanently 
everted annulus. In many plusiines, this basal sclerotization has been reduced to two lateral bars, one 
(commonly the LHS) or both of which are also frequently absent. In the Argyrogrammatini , the RHS 
bar represents the only basal sclerotization of the vesica and is inverted when the vesica is inverted 
and retracted. Further, this bar has heavier dorsal and ventral sclerotization, which gives it a grooved 
appearance, and in extreme instances (e.g. Plusiopalpa), the RHS bar appears to be longitudinally 
split. Some other species also have a RHS bar as the only basal sclerotization (e.g. Rachiplusia) but in 
these taxa, the bar is uniformly sclerotized. In Zonoplusia, the sclerotization is reduced to two, short, 
rudimentary bars, while the dorsal part is incomplete basally in Eutheiaplusia. The bar is very thin in 
Stigmoplusia but the differential sclerotization can be discerned in the somewhat spatulate apex. 

174 Basal RHS lateral sclerotization of vesica produced and turned dorsally to form a smooth, convex, 
rounded plate (Figs 397, 401) (\)lnot (0). 

175 Sub-basal RHS auricular cornutus present (Figs 383, 384, 385) (l)labsent (0). 

Female abdomen Al-7 

176 Pleural membrane of A7 thin, as on A2-6 (0)/ 'pleural membrane of A7 thickened, white, apparently 
glandular and often produced into a shallow pocket (1). The A7 pleural membrane in Puriplusia is 
only slightly thickened and apparently glandular. The thickening in Allagrapha is restricted to a 
shallow pouch on the posterior margin of A7. Autoplusia possesses a shallow pouch also, which is 
produced into a deep, conical invagination in 'Syngrapha' gammoides. Loboplusia has a pleuron 7 
that appears to be glandular but is not thickened. It is, however, difficult to be certain, so the genus 
was coded 0. In one of the two specimens of Brachionycha, pleuron 7 is thickened and white but is not 
apparently glandular. Nevertheless, the genus was coded 1. 

177 Lateral edges of T7 convex or ± straight (Fig. 224) (0)/concave (Fig. 225) (1). This character is almost 
certainly correlated with the last. The lateral margin of T7 in Allagrapha is only slightly concave 
(probably the result of the thickened area of pleuron 7 moving to the posterior margin of the 
segment), while in Puriplusia, the margin is straight (probably due to the pleuron in this genus being 
very slightly modified). The lateral margins of T7 in Loboplusia appear concave and the genus was 
scored 1, although the condition is somewhat uncertain. If the lateral margins are concave, and 
characters 176 and 177 are correlated, then it would be predicted that the examination of fresh 
specimens of Loboplusia would reveal, at least, a glandular, slightly thickened pleuron 7. 

178 Post-spiracular bar and anterior apodeme of segment St2 unmodified, pleural pouch absent (Fig. 234) 
(0)/post-spiracular bar and anterior apodeme of St2 modified to support a large pleural pouch (Figs 
230, 231, 233) (l)/as (1) but pleural pouch small and partially concealed by the tympanal hood (Fig. 
232) (2). This character was frequently unobservable in many specimens due to loss of the relevant 
parts on removal of the abdomen or during subsequent preparation. Adeva, Polychrysia, Eosphor- 
opteryx, Chrysanympha, Lophoplusia and Paectes were scored, by necessity, from male specimens. 
Chalcopasta lacks a post-spiracular bar. 

Anal papillae & A8-A9 + 10 intersegmental membrane 

179 Anal papillae ± square or tapered apically, inner and outer surfaces parallel in dorso-ventral plane 
(Fig. 414) (0)/ 'anal papillae flared apically , inner surface partially or fully facing posteriorly (Figs 417, 
418) (1). 

180 Inner surface of anal papillae membranous and devoid of setae (except for the extreme apex) (Fig. 414) 
(0)/ inner surface densely setose (Figs 417, 418) (1). 

181 Apex of anal papillae entire (Figs 414, 417) (0)1 apex with fine, irregular, setose prominences, giving a 
frayed appearance (Fig. 418) (1). 

182 Anal papillae setae long, fine and undifferentiated (Fig. 420) (0)/ 'anal papillae with a dorso-ventral line 
of strong, pale-socketed setae, one-third to half way from the base, remaining setae mostly small and 
fine (Figs 414, 415, 416) (1). Oncocnemis has a central area with very stout, bluntly pointed setae 
sub-basally and a subapical row of similar but recurved setae. However, this setal type is autapomor- 
phic for the genus and, because the basal and apical setae are of the long, fine type found in the other 
outgroup genera, Oncocnemis was coded 0. 

183 Anal papillae ± square, longer than wide or highly elongate (Figs 414, 415, 416) (0)1 anal papillae short, 
distinctly wider than long (Fig. 419) (1). The plesiomorphic condition encompasses all anal papilla 
forms from the square, somewhat apically lobed type of most plusiines to the highly elongate, 
pointed type of Brachionycha and Chalcopasta. The form found in the euteliines and stictopterines 



SYSTEMATICS OF THE PLUSIINAE 109 

has been coded separately above (character 179) and the genera of these subfamilies are coded * for 
character 183. 

184 Membrane between A8 and anal papillae dorsally extended as a pouch (Fig. 412) (1)/ membrane not so 
modified (Fig. 413) (0). The presence of the pouch has the effect, when everted and extended, of 
turning the anal papillae ventrally. In addition, the pouch may be strongly evaginated but this state 
proved too difficult to score consistently and was not employed. 

185 Base of posterior apophysis dorsally membranous or with a sclerotized flange indistinctly differentiated 
from the base of the anal papilla (Figs 416, 420) (0)/ 'dorsal flange of posterior apophysis sub-basal, 
distinctly separated from the base of the anal papilla and often elongate and leaf-like (Figs 414, 415) (1). 
Many plusiines possess a basal flange on the posterior apophysis but only in the Argyrogrammatini is 
it distinct from the base of the anal papilla. In other genera (e.g. Allagrapha, Puriplusia, Lophoplu- 
sia, Anagrapha), the flange is an irregular sclerotized patch in the angle between the anal papilla and 
the apophysis or on the apophysis but connected to the base of the anal papilla by a semi-sclerotized 
zone of granular membrane. 

186 Membrane between lamina post-vaginalis (LPV) and anal papillae unmodified, extension of anal 
papillae ventrally easy (Q)l membrane between LPV and anal papillae short, thickened and invaginated, 
preventing protraction of anal papillae ventrally (1). 

A8, ostium bursae & antrum 

187 T8 uniformly sclerotized laterally (Fig. 424) (0)/T8 with a subdorsal, longitudinal, desclerotized band, 
the ventral edge of which is continuous with the dorsal edge of the anterior apophysis (Figs 422, 423) 
(1). Chalcopasta also has a desclerotized band but this has its apex along the midline of the anterior 
apophysis, so that the ventral edge of the desclerotization is ventral to, not continuous with, the 
dorsal edge of the anterior apophysis. In Lophoptera and the euteliines, the anterior apophysis is 
absent (see 190). All of these genera were coded 0. 

188 Anterior edge of T8 ventral to anterior apophysis flat (Fig. 424) (0)/ 'invaginated as a dorso-ventral 
groove, with a ventral pocket (Figs 421 , 422, 423) (1). The ventral pocket is very small in Mouralia but 
is enlarged, inflated and somewhat recurved in Abrostola. 

189 T8 irregularly cut back antero-subventrally below the origin of anterior apophysis, leaving the ventral 
base of anterior apophysis 'free' (Fig. 425) (l)/T8 not cut back antero-subventrally, base of anterior 
apophysis ventrally continuous with anterior edge ofT8 (0). State 1 for this character is variable in its 
expression. Antoculeora was coded although its genitalia are so highly modified as to render the 
possible expression of state 1 very difficult. 

190 Anterior apophysis present (Figs 422-425) (0)/absent (Figs 426, 429) (1). 

191 Ventral margin of antrum not evaginated and projecting beyond the posterior edge of intersegmental 
membrane A7-A8 (Fig. 439) (0)/ ventral margin of antrum produced as an evaginated spinose or 
heavily sclerotized projection over the ostium bursae (Figs 427, 428, 435) (1). This probably 
corresponds to the 'classic' plusiine character of 'ostium bursae unprotected/protected'. Many, 
probably convergently derived, conditions are included under the apomorphic state. In Allagrapha, 
Diachrysia and Brachionycha, the evagination is poorly sclerotized and membranous. In contrast, in 
Palaeographa and Caloplusia, the evagination is very strongly sclerotized and the intersegmental 
membrane is fused to the projection ventrally. Autoplusia is coded as A. egena exhibits the 
plesiomorphic state. However, 'Syngrapha' gammoides has a very weakly developed apomorphic 
condition, similar to that found in Antoculeora. The state in Loboplusia is suspected to be but it is 
difficult to say as most of the membrane anterior to the lip of the antrum has been removed in the 
preparation of the slide. In Puriplusia and Macdunnoughia, the evagination is limited to the 
subventral area. 

192 Lateral edges of antrum unmodified (0)/ 'produced as two ± spinose pads lateral to the ostium bursae 
(Fig. 431) (\)l further fused with similar projections from the dorso-lateral corners of the antrum to 
form recurved, spinose grooves (Fig. 432) (2). Many of the genera with broad antra (196: 1), especially 
among the outgroups, have somewhat similar lateral grooves to those found in Puriplusia and 
Macdunnoughia but are not heavily spinose or as deeply invaginated. 

193 Ventral margin of antrum and intersegmental membrane A7-A8 ventrally with short, dense, upright 
scales (Fig. 433) (l)/without (0). The entire female genitalia in Diloba are surrounded by a dense pad 
of upright, black scales. These, however, occur primarily on T7 and St7 and are considered 
non-homologous to the upright antrum-mouth scales found in Allagrapha and other genera. 

194 * Antrum without setae (0)/antrum with 1-3 setae subventrally or laterally (Fig. 435) (\)lantrum with 

several setae along inner, ventral edge (Fig. 434) (2). Brachionycha has numerous setae lateral and 
subventral to the antrum but, as these are a continuation of the generally setose condition of T8 and 



110 IAN J. KITCHING 

not a discrete patch, Brachionycha was coded 0. The setae along the inner, ventral edge of the antrum 
in Diloba also form a continuation of those found along the posterior margin of A8 but the genus was 
coded 2 because of their localized occurrence . However, it remains possible that the condition found 
in Diloba represents a convergent development from the condition found in Brachionycha rather 
than a homologue of that found in Pseudeva and Plusidia. 

195 Anterior edge of T8 ± level with ostium bursae (0)/T8 ventrally produced anteriorly to lie along the 
antrum dorsally and exceeding it (Figs 426, 429) (1). This extension of T8 antero-ventrally may 
represent a highly modified anterior apophysis (cf. character 190). This is suggested by the extension 
bearing an apical, dorsal small lobe. However, Anuga has a lateral anterior bump on the margin of T8 
in the position often occupied by the anterior apophysis in other genera. Therefore, until the 
Stictopterinae and Euteliinae are examined in more detail, the present interpretation will be adhered 
to. In Lophoptera, the extensions are apically highly modified into a pair of spinose pads. See also 
character 202. 

196 Antrum a distinctly invaginated, often heavily sclerotized, structure (Fig. 439) (0)/antrum a broad, 
shallow, open bowl, ostium bursae somewhat exposed (Fig. 434) (1). Brachionycha and Stictoptera 
have narrow, shallow antra and were coded 0. Diachrysia, Allagrapha, Puriplusia and Macdun- 
noughia have invaginated but still somewhat open antra with marginal folds (see character 191) that 
protect the ostium bursae. These genera were coded 0. 

197 Antrum at least with sclerotized granulations, often very heavily sclerotized, well differentiated from 
ductus bursae (0)/antrum membranous, barely differentiated from the ductus bursae (1). 

198 * Antrum sclerotization (if present) free from T8 or continuous only at the antero-ventral corner (Fig. 

434) (0)/antrum with two subdorsal, apically broad and truncate bars, T8 ventral edges extended 
towards the midline (Fig. 433) (l)lextended T8 and antrum subdorsal bars fused, membranous area 
between the ventral edges of T8 very narrow (Fig. 437) (2)/ventral edges of T8 fused; T8, LPV and 
antrum forming a single, sclerotized area (Fig. 428) (3). Mouralia shows a state similar to state 2 but 
the T8 edges are not parallel and have not been extended (as indicated by the presence, in this genus 
of setae along the inner, ventral edge of T8). Setae are absent along the ventral edge of T8 in taxa 
showing states 1-3 (but, it should be noted, such setae are also absent in several genera showing state 
0; e.g. Omorphina, Diachrysia, Allagrapha, Syngrapha). Pseudoplusia has extended edges to T8 but 
no subdorsal bars from the antrum. Autoplusia is coded 2 (the state shown by both A. egena and A. 
olivacea) but 'Syngrapha' gammoides displays state 3. 

199 Anterior apophysis arising approximately halfway down T8 in lateral view (Fig. 430) (0)/anterior 
apophysis arising approximately level with the ventral edge of T8 (Fig. 440) (1). Taxa lacking apodeme 
8 (character state 190:1) were coded *. 

200 Antrum large, heavily sclerotized, smooth, open and curved somewhat to the RHS (Figs 437, 439) 
(1)/ 'antrum relatively small, often poorly sclerotized, with longitudinal folds and ridges, not forming an 
open cavity (0). The antra of Eosphoropteryx and Palaeographa are curved to the LHS. 

201 Membrane lateral and anterior to ostium bursae thin and unmodified (Q)lmembrane thick, white, 
folded and tough (1). The thickened membrane in Allagrapha is the same membrane as that coded in 
character 176 and is thus most probably convergent on that condition found in, for example, 
Palaeographa. The membrane in Euchalcia is only slightly thickened. 

202 T8 dorsally with anterior edge ± level with or anterior to the antrum (0)/7S represented by a thin band 
dorsally, anterior edge posterior to the antrum (Fig. 426) (1). T8 in Adeva (Fig. 430) is reduced to a 
thin band dorsally but this is formed by a strongly cut-back edge from an anteriorly positioned 
anterior apophysis. It is considered to be an independent derivation from state rather than 
homologous to the condition found in the euteliines and Lophoptera. 

Ductus bursae & corpus bursae 

203 Ductus bursae (DB) short (less than half the length of abdomen), usually broad (Fig. 458) (0)/DB long 
(over three-quarters the length of the abdomen) and narrow (Fig. 446) (1)/DB over 2 X as long as 
abdomen and very narrow (Fig. 447) (2). 

204 Long axis of corpus bursae (CB) ± parallel with long axis of DB (Fig. 449) (0)/long axis of CB ± at 
right angles to long axis of DB (Figs 446, 447, 448) (1). 

205 Membrane of CB, at point of entry ofDB, unmodified (0) / 'thickened, white and folded (1). 

206 Median section of DB tubular or merely flattened (Fig. 446) (0)/median section expanded somewhat 
laterally with dorsally recurved edges (Fig. 441) (1). Dactyloplusia has a laterally expanded median 
section of the DB but the edges are not recurved. 

207 DB with a lateral LHS short pouch with thick walls (\)lwithout (0). 



SYSTEMATICS OF THE PLUSIINAE 111 

208 CB relatively inelastic, ± maximum size when dissected (0)/ CB with very elastic walls, very small when 
first dissected but can be very easily stretched to several times its original size (1). 

209 * Ductus seminalis arising from apex ofCB (Figs 450, 461, 462, 463) (0)/ 'from fundus (Fig. 444) (l)/from 

an accessory bursa produced from the DB (Fig. 460) (2). There is considerable difficulty in some taxa 
(e.g. Mouralia, Diachrysia, Puriplusia) in deciding which of two pouches is an enlarged apex and 
which is the CB proper. In Puriplusia, the orientation of the grooves near the origin of the DB 
(character 213) suggests that there may have been a 180 degree change in the orientation of the CB so 
that the apex is now directed anteriorly. However, this is not certain and pending further work, 
Puriplusia is coded 1. The same situation may pertain to Loboplusia. In Sclerogenia, the apex is 
interpreted to be recurved and totally fused with and open to the CB. Erythroplasia (Fig. 453) is also 
very problematical. A fundular origin of the ductus seminalis is inferred but it could be that the CB is 
extremely reduced and the apex very elongate to give the observed 'boomerang-shape', resulting in 
the ductus seminalis then being interpreted as arising from the apex and changing the coding for the 
genus to 0. The second species of Erythroplusia, E. pyropia, is very similar to E. rutulifrons and is of 
no use in resolving this problem. Those genera exhibiting state 1 for character 207 also present 
difficulties. The ductus seminalis is interpreted as arising from the apex in Trichoplusia and 
Thysanoplusia, and from the fundus in Zonoplusia and Eutheiaplusia, but this is by no means certain. 
Plusiopalpa is interpreted as having a small, recurved apex, bearing the ductus seminalis, with a 
secondary large apical pouch present dorsal and posterior to it. The ductus seminalis in Pseudoplusia 
is considered to arise from a long, anteriorly-recurved apex that is slightly fused to the CB basally (see 
character 214). Chrysodeixis is interpreted as having a fundular origin but examination of other 
species in this genus (C. acuta, [Dufay, 1970a]; C. illuminata, BM noctuid slide 11649) shows a similar 
condition to that found in Pseudoplusia, but in which the origin of the recurved apex has moved to a 
position about halfway along the CB. The extreme fundular origin of the ductus seminalis in C. 
chalcites may merely represent the ultimate expression of this transformation. The origin of the CB 
from the DB in Stictoptera is less well defined than in most species but the ductus seminalis still 
appears to arise from the apex of the CB, rather than a lateral origin on the DB. A similar 
interpretation was followed for Magusa, Paectes and Anuga, although the condition found in 
Lophoptera casts some doubt on this interpretation. More work is evidently needed in the 
stictopterine-euteliine complex. 

210 *CB with weakly defined and irregular folds and lines, often smooth (Fig. 463) (0)/with regular, 

elongate, longitudinal ridges, originating from dorsal and lateral longitudinal 'focal' ridges, which may 
or may not be sclerotized (Fig. 464) (\)l ridges forming a ± circular pattern (Fig. 457) (2). 

211 CB without a localized signum (0)/CB with a small, slightly concave, circular signum (Fig. 457) 
{\)lsignum invaginated as a narrow internal ridge (Fig. 465) (2). 

212 CB with uniform internal ornamentation or, if several forms present, then these gradually merge one 
into another (0)/CB with a distinct discontinuity halfway along its length, separating a pale, membra- 
nous posterior half from an anterior half covered in sclerotized granulations and finer folding (Figs 
454, 456) (1). 

213 * Area of entry of D B into CB with large, strong, well sclerotized internal folds that extend onto the apex 

or rarely onto the fundus (Figs 450, 451, 454, 455, 456) (3)/ridges present and sclerotized but not 
strongly developed (Figs 445, 449) (2)lridges not sclerotized, merely granular (Fig. 452) (\)lridges 
absent (Fig. 461) (0). Calophasia has a smooth area of sclerotization in the dorsal angle between the 
DB and the apex, that may have a few folds. However, these folds are not extensive and the genus 
was coded 0. Paectes has a long series of diagonal, heavily sclerotized folds on the posterior section of 
the CB but these are not connected with folds arising on the DB and are not considered homologous 
to the folds found in Autoplusia and related genera. The apex and folds are highly reduced in 
Allagrapha, while in Puriplusia, they continue onto the fundus rather than the apex (but see 209). 

214 Apex recurved in dorso-ventral plane and somewhat fused basally to the CB (Figs 443, 444) (\)lapex 
not so modified (0). 

215 CB without a dorso-lateral RHS patch of thickened white membrane somewhat anterior of the 
entry-point of the DB (0)/such a patch present (Figs 461 , 462) (1)/ 'patch expanded into a shallow pouch 
(Fig. 463) {2)1 expanded further into a large pouch or a long appendix in which the thickened membrane 
is present only apically (Fig. 458) (3). 

Miscellaneous 

216 Colour pattern monomorphic Wpattern sexually dimorphic, males greyish, females brown (Figs 82, 
83) (1). In Autoplusia, only 'Syngrapha' gammoides is obviously sexually dimorphic in this manner. 
The difference is only barely discernible in other species of the genus (A. egena, A. olivacea). In 



112 IAN J. KITCHING 

Sclerogenia, the females appear to be slightly duller than the males but this is not great enough to 
warrant the description 'sexually dimorphic'. Cucullia is also sexually dimorphic but shows a 
different type of variation (males have white hindwings, females brown) and was coded 0. Stictoptera 
is highly polymorphic but the various forms are not sex limited. 

Analytical procedure 

The data were analysed using David Swofford's program for inferring phylogenies using the principle of 
maximum parsimony, PAUP (version 2.2), run on the IBM System/370 3032 at the Rutherford Appleton 
Laboratory. Due to the large size of the data set (57 taxa by 216 characters), the analysis was broken down 
into several subprocedures that were run separately at different times. 

Initially, a simple Wagner tree was constructed using the addition sequence CLOSEST and setting 
HOLD (the number of trees held in memory at any one step of tree construction) equal to one. No 
branch-swapping was employed at this point. The result was a single cladogram of 659 steps and a 
consistency index (Kluge & Farris, 1969) of 0-466. 

This cladogram was subsequently used as an input to a branch-swapping analysis (using the option 
SWAP = ALT in an attempt to reduce the run time), which also invoked the MULPARS option to check 
for the existence of multiple, equally-parsimonious cladograms. A default maximum of 50 cladograms was 
found, with the reduced length of 648 steps and a consistency index of 0-474. 

These 50 cladograms were then evaluated in order to determine the topological differences between 
them. It was found that they consisted of five pairs, one triplet and one quadruplet of mutually-exclusive 
subcladograms and a single alternative that was mutually exclusive to all but one of the five pairs. 
Combining these alternative subcladograms gives a minimum number of 448 equally-parsimonious 
cladograms for this data set. 

In order to determine which of these 448 possibilities represented the best hypothesis of the interrela- 
tionships of the plusiine genera, the apomorphy lists for six (output cladogram numbers 27, 29, 39, 44, 45 
and 46) were obtained, which covered all the alternative subtopologies. A single preferred cladogram was 
then identified by a process of character analysis and evaluation, which is described in the next section. 

A second analysis was then performed on a restricted data set, with the outgroup taxa (Chalcopasta- 
Anuga, 48-58) excluded. The intention was to check whether the 64 alternative topologies found for the 
plusiines alone were exhaustive or whether, by restricting MAXTREE (the maximum number of 
equally-parsimonious trees held in memory during a branch-swapping procedure) to 50, any had been 
missed. To save computing time , the preferred topology (see below) was entered as the initial user-tree and 
branch-swapping performed using SWAP = ALT, MULPARS and MAXTREE = 50. The initial 
cladogram of 451 steps (it is worth noting in passing that 197 or 30-6% of the steps on the original cladogram 
occurred outside the Plusiinae) was, surprisingly, reduced in length by two steps to 449 (consistency index 
not calculated) and only 16 alternatives were found. The two missing pairs were those involving the 
location of Loboplusia and the sequence of branching among the various tribes and subtribes (see below) . 

Initially, it was suspected that this additional shortening of the cladogram was an artefact caused by the 
omission of the outgroup taxa. Thus, a final analysis was undertaken in which one of the previously 
obtained outgroup topologies was added to one of the 16 plusiine topologies and the resultant cladogram 
entered as a user-tree. 

Fitting of the data to this cladogram resulted in a length of 647, a reduction of one step. Thus the 50( + ) 
cladograms of 648 steps obtained originally did not represent the minimum-length as was first thought. The 
branch-swapping algorithm applied in the first analysis had failed to find the minimum-length topology 
before the MAXTREE limit of 50 had been reached. Further, it is unlikely that it would have been found 
even had MAXTREE been set to the system maximum of 150, given a minimum of 448 possible 
cladograms of 648 steps and the basal position of the swap necessary to locate the shorter cladograms of 647 
steps. (The algorithm appears to perform its branch-swapping starting with pairs of terminal taxa and 
proceeding towards the base of the cladogram. Also, to find the 647-step cladograms, a swap involving the 
rearrangement of subcladograms of 8 or more taxa was required, based upon an initial topology that 
occurred only five times out of the original 50 cladograms and which was first found at cladogram 33 . ) Had 
the second analysis of the restricted data set not been performed, the existence of the 16 cladograms of 647 
steps would never have been revealed. 

However, examination and evaluation of the apomorphy lists of the 647-step cladograms demonstrated 
that the branching sequence of the major plusiine lineages was determined by very similar character state 
changes to those present on one subset of the rejected 648-step topologies. Thus, the minimum-length 
cladograms were rejected in favour of the 648-step, preferred cladogram referred to above. The full 
reasoning behind this decision is given below. 



SYSTEMATICS OF THE PLUSIINAE 113 

The 648-step cladograms 

1. Agrapha and Plusiopalpa 

Two subcladograms were found in which the positions of Agrapha and Plusiopalpa differed. Subcladogram 
1 (Fig. 9) places Agrapha as the sister-group of Argyrogramma + Stigmoplusia (A + S), and Plusiopalpa as 
the sister-group of Acanthoplusia + Ctenoplusia (A + C). Subcladogram 2 (Fig. 8), in contrast, treats 
Plusiopalpa as the sister-group of A+S, and Agrapha as the sister-group of these three genera plus A + C. 
The different topologies are due to alternative interpretations of 13 characters (61, 79, 83, 86, 87, 88, 92, 
98, 105, 130, 138, 144, 162). An evaluation of these characters is required in order to decide which 
subcladogram represents the best cladistic hypothesis of the relationships between the six genera. 
61: The sequence of suppression of the male fore-tibial spines forms a clear, non-homoplasious 
transformation series on subcladogram 2, where state 1 characterizes A+S + Plusiopalpa, state 2 
unites A +S, and state 3 is autapomorphic for Argyrogramma. Character 61 is not as convincing on 
subcladogram 1 , on which states 1 and 2 are interpreted as independent derivations from state 0. On 
both cladograms, state 1 is also convergently derived in Plusiotricha and in the stictopterines (see 
also below). Thus character 61 supports subcladogram 2. 
79: This character is problematical on both subcladograms. The expected transformation series, in 
which the hair tufts on segments A5 and A6 both gain discrete edges (state 1), followed by 
independent losses of such margins on A6 (state 2) and A5 (state 3), did not appear. On 
subcladogram 1 , A +S + Agrapha are characterized by the change — > 3 (discrete edges on A6 only) , 
followed by independent derivations of state 2 in Stigmoplusia ( A6 only — » A5 only) and state 1 in 
Agrapha ( A6 only — > A5 and A6). On subcladogram 2, state 1 in Agrapha is independently derived 
from state 0, with .4+5 characterized by the change — > 3 followed by the autapomorphic change 
3 — > 2 in Stigmoplusia. Alternative optimizations on subcladogram 2 are equally ambiguous. One 
assigns state to the node immediately ancestral to Argyrogramma and Stigmoplusia, in which case 
states 2 and 3 respectively are independently derived in the two genera from state 0. Alternatively, 
the internal node could be assigned state 2, whence Argyrogramma is characterized by the 
autapomorphic change 2— > 3. Subcladogram 1 is marginally favoured in that character 79 delimits a 
monophyletic group on this subcladogram but the transformation series is not very satisfactory, 
despite a character consistency of 1-000 (true also of subcladogram 2). 
83: This character is one of the most highly homoplasious in the data (consistency = 0-313). However, 
state 4 is only found in Plusiopalpa and Argyrogramma and thus lends weak support to subclado- 
gram 2 (Stigmoplusia possess the widely distributed state 3). 
86: Subventral accessory pockets unite A+S and Agrapha as a monophyletic unit and thus support 
subcladogram 1. On subcladogram 2, these pockets have to be hypothesised to have developed 
independently in the two groups (as well as in Acanthoplusia, which is also true on subcladogram 1). 
87: Although optimization of this character on subcladogram 2 suggests an initial gain followed by a 
'monophyletic' loss in A+S + Plusiopalpa, an equally parsimonious interpretation is one of 
independent gain in A + C and Agrapha. This is also the optimized distribution of states for this 
character on subcladogram 1 and thus character 87 is of no use in deciding which subcladogram 
should be accepted. 
88: These characters show similar distributions to the last, in that they can be interpreted on subclado- 
98 gram 1 as having been independently gained (Agrapha and Argyrogramma) or gained once (thus 
705 characterizing A+S + Agrapha) with subsequent reversal (in Stigmoplusia). The former distribu- 
tion is that found on subcladogram 2 and so characters 88, 98 and 105 are also irrelevant to the 
decision. 
92: On subcladogram 1, the state changes 1 — > 3 and 1 — » 2 arise independently and characterize A+S 
and A + C + Plusiopalpa respectively. On subcladogram 2, state 3 transforms into state 2, resulting 
in the exclusion of Agrapha. Although both subcladograms are equally supported, other shared 
similarities of the secondary hair pencil attachments between A +S and Plusiopalpa (see below) tip 
the balance slightly in favour of subcladogram 2 and its sequential transformation series. 
130: A juxta that is free from the saccular flanges also occurs in Pseudoplusia + Chrysodeixis and thus 

offers only weak support for the monophyly of A+S + Agrapha (subcladogram 1). 
138: Similarly, but conversely, the monophyly of A +S + Plusiopalpa (subcladogram 2) can be weakly 

supported by the monophyletic reversal of this character. 
144: This character (the degree of development of the clavus) shows an extremely high amount of 
homoplasy (character consistency index = 0-133) and thus is very suspect. Both subcladograms 
postulate the reduced condition (state 1) to be plesiomorphic for the six genera. Subcladogram 1 
hypothesises independent reversals to a long clavus (state 0) in Argyrogramma and A + C + 



114 



IAN J. KITCHING 




Argyrogramma 



Stigmoplusia 



Plusiopalpa 



Acanthoplusia 



Ctenoplusia 



Agrapha 



Plusiotricha 
Eutheiaplusia 




Argyrogramma 



Stigmoplusia 



Agrapha 



Plusiopalpa 



Acanthoplusia 



Ctenoplusia 



Plusiotricha 
Eutheiaplusia 




Lamprotes 



Pseudeva 



Plusidia 



Euchalcia 



Adeva 



Panchrysia 
Chrysanympha 




Lamprotes 



Pseudeva 



Plusidia 



Euchalcia 



Adeva 



Panchrysia 
Chrysanympha 



Figs 8-11 8, subcladogram 2, depicting Plusiopalpa as the sister-group of Argyrogramma + Stigmoplusia 
and a relatively basal position for Agrapha. 9, subcladogram 1, illustrating the alternative placements of 
Plusiopalpa and Agrapha. 10, subcladogram 3, showing Plusidia as the sister-group of Lamprotes + 
Pseudeva. 11, subcladogram 4, the alternative position of Plusidia as the sister-group of Euchalcia + 
Adeva. 



SYSTEMATICS OF THE PLUSIINAE 115 

Plusiopalpa, while subcladogram 2 suggests a reversal above the point where Agrapha branches off, 
followed by a re-reversal to state 1 in Stigmoplusia. Both subcladograms include an autapomorphic 
extreme reduction in Agrapha. Given the number of steps associated with this character, it is 
considered to be too unreliable and of little use, especially with regard to the current decision. 
162: A+S + Agrapha are united on subcladogram 1 by the (highly homoplasious) gain of a subapical 
cornutus (SAC). However, examination of species other than the type species shows a SAC to be 
general in Ctenoplusia (Dufay, 1970a). C. limbirena is apparently aberrant in lacking one, so further 
increasing the homoplasy in this character. Thus, absence of a SAC is rejected as demonstrating 
relationship. 
Overall, the decision between the two topologies was close. Subcladogram 1 is supported by character 86 
and weakly by characters 79 and 130. Subcladogram 2 is supported by character 61 and weakly by 
characters 83, 92 and 138. On balance, the evidence in favour of subcladogram 2 appears slightly better and 
it is accepted over subcladogram 1. However, the most expedient solution may be the consensus 
tetrachotomy (see below). 

2. Plusidia 

Two subcladograms were found in which the position of Plusidia was ambiguous. In subcladogram 3 (Fig. 
10), Plusidia is treated as the sister-group of Lamprotes + Pseudeva (L + P), while in subcladogram 4 (Fig. 
11), the genus is placed as the sister-group of Euchalcia + Adeva (E+A). Ten characters (6, 41 , 42, 58, 144, 
154, 171, 186, 194, 205) are involved in the ambiguity. 

Five characters support a sister-group relationship between Plusidia and E+A. Of these, four are 
reversals in highly homoplasious characters (6, 41, 42, 58), while the apomorphic state for the fifth (205) 
also characterizes Cucullia, Trichoplusia and Plusiotricha + Eutheiaplusia. The evidence is thus not very 
strong. In contrast, the five characters that suggest that Plusidia is most closely related to L + P include two 
(171 and 186) that are unique and unreversed. Further, the transformation series of character 194 is only 
homoplasious insofar as state 1 is independently derived in the outgroup taxon Diloba, and there is doubt 
as to the homology of this condition and that found in Plusidia + L + P (see below). Character 154 is found 
in its apomorphic state in Panchrysia, Chrysanympha and E+A and thus is interpreted as reversed in 
Plusidia + L + P on subcladogram 3, while the last, character 144, is one of those for which extreme doubt 
was expressed in the previous section regarding reliability. 

Overall, there can be little doubt, after evaluating the characters supporting the alternative subclado- 
grams, that subcladogram 3 is to be preferred and Plusidia interpreted as the sister-group of Lamprotes + 
Pseudeva. 

3. Autographa, Cornutiplusia and Plusia 

It was noted during the data collection that few synapomorphies were being found for Autographa, 
Cornutiplusia, Plusia and their immediate relatives. Thus, it was not unexpected when multiple, equally- 
parsimonious subcladograms were found that included these genera. In all of these subcladograms, there 
was a constant terminal cluster of four genera (Caloplusia, Syngrapha, Palaeographa, Lophoplusia; = 
CSPL) and a constant outgroup, Anagrapha. Between these taxa, four topologies were found that 
rearranged Plusia, Cornutiplusia, Autographa gamma and A. bractea. 

Three subcladograms, 5, 6 and 7 (Figs 12-14 respectively) agree in placing A. gamma as the basal taxon, 
while subcladogram 8 (Fig. 15) so places A. bractea. Three subcladograms, 5, 7 and 8, also agree in placing 
Plusia as the sister-group of CSPL, while subcladogram 6 places Cornutiplusia in this position. Subclado- 
gram 5 differs from subcladogram 7 in placing Cornutiplusia and A. bractea as sister taxa. Four characters 
only are involved in differentiating the alternative topologies: 12, 64, 166 and 215. 

Character 12 is highly homoplasious (consistency index = 0T00). The apomorphic state varies from a 
mere silver spot between the reniform stigma and the cubital vein (as seen in Antoculeora and Plusiopalpa) 
to that in which the reniform stigma is almost entirely outlined in silver (a condition unique to Lophoplusia 
and perhaps not too far removed from the completely silver stigmata of Plusia and A. bractea). However, 
character 12 is one of the two characters that support a sister-group relationship on subcladogram 6 
between Cornutiplusia and CSPL. The other character on this branch, character 64, is the 'classic' plusiine 
character of spined hind tibiae. This is also highly homoplasious; state 1 is also found in Autoplusia and 
state 2 in Rachiplusia and Anagrapha. The topology of subcladogram 6 is also supported by the observation 
that occasional spines occur in Plusia and Autographa (McDunnough, 1944). 

Plusia is placed as the sister-group to CSPL on subcladograms 5, 7 and 8 by a synapomorphy in character 
166. State 2, the distinctly winged subapical cornutus, was scored as autapomorphic for Lophoplusia and as 
'not comparable' for Palaeographa, Syngrapha and Caloplusia, because the type species of these taxa lack 
subapical cornuti. That these three taxa are predicted to show state 2 is interesting. Several species of 



116 



IAN J. KITCHING 




Caloplusia 
Syngrapha 
Palaeographa 
Lophoplusia 



Plusia 



Autographa bractea 



Cornutiplusia 



Autographa gamma 



Anagrapha 




Caloplusia 
Syngrapha 
Palaeographa 
Lophoplusia 



Cornutiplusia 



Plusia 



Autographa bractea 



Autographa gamma 



Anagrapha 




Caloplusia 
Syngrapha 
Palaeographa 
Lophoplusia 



Plusia 



Cornutiplusia 



Autographa bractea 



Autographa gamma 



Anagrapha 




Caloplusia 
Syngrapha 
Palaeographa 
Lophoplusia 



Plusia 



Cornutiplusia 



Autographa gamma 



Autographa bractea 



Anagrapha 



Figs 12-15 The four alternative subtopologies involving Autographa, Plusia and Cornutiplusia. 12, 
subcladogram 5. 13, subcladogram 6. 14, subcladogram 7. 15, subcladogram 8. 



SYSTEMATICS OF THE PLUSIINAE 117 

Syngrapha, particularly those considered by Eichlin & Cunningham (1978) to be primitive (e.g. S. 
ottolenguii (Dyar), 5. viridisigma (Grote), 5. orophila (Hampson)), do bear subapical cornuti on the vesica 
and in one species, 5. epigaea (Grote), the cornutus even appears to be winged (Eichlin & Cunningham, 
1978: fig. 58a). Thus, the interpretation of Plusia as the sister-group of CSPL, on the basis of character 
state 166:1, may prove well-founded and is provisionally accepted here as the best cladistic hypothesis. 
Subcladogram 6 is thus rejected, despite the evidence furnished by character 64. 

A. gamma is placed as the most basal taxon on subcladograms 5-7 because it displays the plesiomorphic 
state for character 215. The first apomorphic state is the presence of a dorso-lateral patch of thickened 
membrane on the corpus bursae and is found in Plusia and Lophoplusia. This is further elaborated into a 
shallow pouch (state 2) in A. bractea and into a large pouch or appendix (state 3) in Cornutiplusia. All 
subcladograms require a reversal from state 1 to state in Palaeographa + Syngrapha + Caloplusia. 
Subcladogram 7 has what appears to be the worst optimization of states, requiring the derivation of state 2 
directly from state 0, then a further autapomorphic development of state 3 in Cornutiplusia, together with a 
reduction in the structure, stage 2 — > 1, along the branch leading to Plusia + CSPL. Subcladogram 8 is 
somewhat better in that independent derivations of states 2 and 3 from state are postulated, a distribution 
of states also found on the rejected subcladogram 6. Subcladogram 5, however, unites A. bractea and 
Cornutiplusia using state 2 and then derives state 3 autapomorphically in the latter taxon. This interpreta- 
tion is further supported by the observation that state 2 appears to be present in all the putative North 
American relatives of A. bractea except one, as judged from the figures and phylogenetic tree given by 
Eichlin & Cunningham (1978) (but see Clade analysis). Thus, the branching sequence, and associated 
character state changes, of subcladogram 5 is accepted as the best cladistic hypothesis of relationships for 
these genera. 

4. Loboplusia 

The genus Loboplusia, with its large amount of missing data and relatively simple male genitalia, was 
expected to be ambiguous in position. Two subcladograms were found; subcladogram 9 (Fig. 16) places 
Loboplusia as the sister-group of Erythroplusia, while subcladogram 10 (Fig. 17) hypothesises a sister- 
group relationship between Loboplusia and Autoplusia + Antoculeora + Sclerogenia (AAS). Eleven 
characters (6, 12, 59, 113, 142, 148, 155, 176, 177, 209) are involved in these alternative interpretations of 
the position of Loboplusia. 

Loboplusia is grouped with Erythroplusia on the basis of three character state changes. The loss of the 
scale tooth on the forewing tornus (6) is highly homoplasious and of little importance. Similarly, the 
fundular origin of the ductus seminalis on the corpus bursae (209) appears to have been multiply derived, 
supporting the reservations outlined above regarding the correct interpretation of corpus bursae morphol- 
ogy. Finally, character 155 was scored as 'missing data' for Loboplusia, because the transtillae were torn. 
Thus, the presence of the apomorphic state of character 155 in Loboplusia is a prediction of the 
optimization procedure and not an empirical observation. The evidence supporting subcladogram 9 is 
therefore tenuous. 

On subcladogram 10, Loboplusia is placed by four characters in a larger clade including AAS and a clade 
consisting of Allagrapha, Macdunnoughia and Puriplusia (AMP). Two of these characters, 142 and 176, 
are subsequently reversed on the branch subtending Loboplusia. The condition of character 177 was 
considered somewhat uncertain but does provide weak support for this topology. The apomorphic state of 
character 113 is shared by Loboplusia and AMP, and is considered to have reversed in AAS. Further, 
Loboplusia is placed as the sister-group of AAS by three characters, all with low consistency indices. Two 
(59 and 148) are reversals that occur several times elsewhere on the cladogram, while the third (12) was 
rejected above as unreliable. 

Overall, the evidence supporting each placement of Loboplusia is very weak, with subcladogram 10 
being marginally preferable. However, until such time as further studies are performed and additional 
specimens become available, the best course may be to treat Loboplusia as forming a trichotomy with 
Erythroplusia and AAS + AMP (see below). 

5. Rachiplusia and the branching sequence of the tribes and subtribes 

The final pair of alternative topologies within the Plusiinae affects the fundamental branching pattern of 
the Argyrogrammatini and the three subtribes of the Plusiini: the Plusiina, Euchalciina and Autoplusiina. 
(The Autoplusiina are temporarily defined as Rachiplusia, Erythroplusia, Loboplusia, AAS and AMP. ) In 
addition, there is ambiguity concerning the relationships of Rachiplusia. 

Subcladogram 11 (Fig. 18) agrees well with the trees put forward previously by Ichinose (19626) (Fig. 2) 
and Eichlin & Cunningham (1978) (Fig. 3). Both the Argyrogrammatini and Plusiini, as currently 
recognized, are monophyletic. Within the latter, the subtribe Euchalciina is the sister-group of the Plusiina 



118 



IAN J. KITCHING 



Autoplusia 



Antoculeora 




Erythroplusia 
IO • Rachiplusia 




Autoplusia 



Antoculeora 



Sclerogenia 



Loboplusia 



Allagrapha 



Macdurmoughia 



Puriplusia 



Erythroplusia 



Rachiplusia 




PLUS UNA 



AUTOPLUSIINA 



Rachiplusia 



EUCHALCIINA 



ARGYROGRAMMATINI 



ABROSTOLINI 




ARGYROGRAMMATINI 



Rachiplusia 



AUTOPLUSIINA 



PLUSIINA 



EUCHALCIINA 



ABROSTOLINI 



Figs 16-19 16, subcladogram 9, showing Loboplusia as the sister-group of Erythroplusia. 17, subclado- 
gram 10, with Loboplusia as the sister-group of Sclerogenia + Antoculeora + Autoplusia. 18, subclado- 
gram 11, the topology of the tribal and subtribal branching that places Rachiplusia as the sister-group of 
the remaining Autoplusiina. 19, subcladogram 12, the alternative subtribal topology illustrating a 
paraphyletic Plusiini and with Rachiplusia as the sister-group of the Argyrogrammatini. 



SYSTEMATICS OF THE PLUSIINAE 119 

+ Autoplusiina. The genus Rachiplusia is placed as the most basal branch within the last of these subtribes. 

The monophyly of the Plusiini is based upon three characters. Two of these (121 and 127) are moderately 
homoplasious but the last, character 1 17, is uniquely derived but reversed in Antoculeora. In this genus, the 
broad lateral flange of the pleurite is absent, a condition probably correlated with the autapomorphic form 
of the pleurite coded by character state 118:2. The sister-group relationship between the Plusiina and the 
Autoplusiina is based upon a single character, 116, the autapomorphic state of which is lost in Antoculeora 
(cf. character 117) and independently derived in Anadevidia. Rachiplusia is placed as the sister-group of 
the rest of the Autoplusiina based on reversals in three highly homoplasious characters (18,59 and 148) and 
a change from state to state 2 in character 213. Although this character is homoplasious (particularly 
when the outgroups are taken into consideration), further work may justify it and the position of 
Rachiplusia it supports. 

The alternative subcladogram 12 (Fig. 19) is in considerable disagreement, in that the Plusiini is 
paraphyletic, not only with respect to the subtribal arrangement, but also because Rachiplusia is treated as 
the sister-group of the Argyrogrammatini. However, the supporting characters are very weak. The 
Argyrogrammatini , Autoplusiina and Plusiina are grouped together to the exclusion of the Euchalciina by 
two characters. The first, 116, was synapomorphic for the Autoplusiina and Plusiina on subcladogram 11, 
on which the presence of state 1 in Anadevidia (the basalmost member of the Argyrogrammatini) was 
interpreted as convergent. On subcladogram 12, state 1 is interpreted as uniquely derived but reversed in 
the Argyrogrammatini above Anadevidia. The second character state change is 83:0— > 3 and concerns the 
degree of development of the sternal hair pencils in the males. The sequence of state changes for character 
83 is very complex and highly homoplasious. Very little confidence can be placed in it. The Autoplusiina 
and Argyrogrammatini are united by two reversals in characters 18 and 148, both of which are subject to 
multiple convergent gains and reversals. Finally, two character state changes support the sister-group 
relationship between the Argyrogrammatini and Rachiplusia: 14:0 — > 2 and 110. The latter (whether the 
vinculum is V-shaped or Y-shaped) is trivial, homoplasious and can be discounted. 14 can also be rejected 
once its full transformation on subcladogram 12 is analysed. The complete series consists of — * 2 
{Rachiplusia + Argyrogrammatini), 2 — ► 1 {Anadevidia only) and two convergent — » 2 changes 
{Autoplusia, Mouralia). The first two of these imply that seta 1 of the female frenulum was first lost and 
then partially regained as a rudiment in Anadevidia. The more parsimonious sequence morphologically, in 
which the seta is reduced and then lost, is precluded by the position of Rachiplusia. However, both 
interpretations for the origin of state 1 in Anadevidia (reduction as an intermediate step or as a terminal 
reversal) are allowed on subcladogram 11. Thus, on the basis of this, and on the evidence provided by 
characters 116, 117 and 213, subcladogram 12 is rejected and subcladogram 11 accepted as the best cladistic 
hypothesis for this section of the plusiine cladogram. 

6. Outgroups 

Due to the variety and 'taxonomic distance' between the outgroups (11 genera from four subfamilies), it 
was not surprising to find that there were many alternative topologies for them. Three constant features 
were found: 

(a) Oncocnemis and Chalcopasta are sister-groups; 

(b) Brachionycha and Diloba are sister-groups; and 

(c) the Stictopterinae and Euteliinae are each monophyletic and together form a monophyletic group to 
which Magusa is the outgroup (= SEM). 

However, the sequence in which these three groups and the other two genera, Cucullia and Calophasia, 
clustered proved very variable. One pair of alternative topologies was found in which Calophasia + 
Cucullia + SEM formed a clade. The first (Fig. 20) places Calophasia as the sister-group of SEM; the 
second (Fig. 21) places it as the sister-group of Cucullia. Accepting this as a terminal taxon, three further 
topologies were found in which its sister-group was Brachionycha + Diloba (Fig. 22), Oncocnemis + 
Chalcopasta (Fig. 23), or these four genera together (Fig. 24). In addition, a non-exclusive alternative 
topology was found (Fig. 25) in which Calophasia + Cucullia was the sister-group of Oncocnemis + 
Chalcopasta, and SEM was the most basal clade. All these relationships have to be qualified by the 
statement that there is no guarantee that the root of the entire cladogram lies between the Plusiinae and the 
outgroups. 

The precise resolution of the relationships of the outgroups is of secondary importance in this study. 
Thus, the decision as to which of the above alternatives is to be preferred is not critical. The topology that is 
depicted on the final preferred cladogram (see below) is that which places the Plusiinae nearest to the 
Stictopterinae, their erstwhile closest relatives, in order to evaluate the characters that support this 
relationship. 



120 



IAN J. KITCHING 




Stictopterinae 
Euteliinae 



Magusa 



Calophasia 



Cucullia 




Stictopterinae 
Euteliinae 



Magusa 



Calophasia 



Cucullia 



Stictopterinae 

Euteliinae 

Magusa 

Calophasia 

Cucullia 





Stictopterinae 

Euteliinae 

Magusa 

Calophasia 

Cucullia 

Oncocnemis 

Chalcopasta 

Brachionycha 

Diloba 




Stictopterinae 

Euteliinae 

Magusa 

Calophasia 

Cucullia 



Brachionycha 
Diloba 
Oncocnemis 
Chalcopasta 




Stictopterinae 

Euteliinae 

Magusa 

Brachionycha 

Diloba 

Oncocnemis 

Chalcopasta 

Calophasia 

Cucullia 



Figs 20-25 The alternative outgroup subtopologies. 20, outgroup subtopology 1. 21, subtopology 2. 22, 
subtopology 3. 23, subtopology 4. 24, subtopology 5. 25, subtopology 6. Note that subtopology 6 is an 
exclusive alternative to subtopologies 1-5. 



SYSTEMATICS OF THE PLUSIINAE 121 

The 647-step cladograms 

Following removal of the outgroups, further global branch-swapping with the multiple parsimony option 
discovered a set of 16 cladograms. After reinstatement of the outgroups, these cladograms were found to 
be 647 steps long, one step shorter than those found in the earlier analyses that were described above. The 
647-step cladograms are of similar topology to the rejected 648-step subcladogram 12, in that Rachiplusia 
forms the sister-group of the Argyrogrammatini, but differ in the branching sequence of the subtribes of 
the Plusiini (Fig. 26). Whereas, on subcladogram 12 of the 648-step cladograms, the Autoplusiina (less 
Rachiplusia) form the sister-group of the Argyrogrammatini + Rachiplusia, on the 647-step cladograms, it 
is the Plusiina that occupy this position. The 16 alternatives arise from the mutually exclusive topologies 
referred to previously as subcladograms 1/2 (Agraphai ' Plusiopalpa) , 3/4 (Plusidia) and 5/6/7/8 (Auto- 
graphal Comutiplusial Plusia) . The presence of further alternatives in the outgroup taxa was not observed 
but all of the topologies found previously would be expected. Reversing the order of branching of the 
Plusiina and Autoplusiina (less Rachiplusia) appears to preclude the alternative topology that places 
Loboplusia as the sister-group to Erythroplasia. 

ARGYROGRAMMATINI 



Rachiplusia 



PLUSIINA 



AUTOPLUSIINA 



EUCHALCIINA 



ABROSTOLINI 

Fig. 26 The branching sequence of the tribes and subtribes on the 647-step cladograms. Note that the 
position of Rachiplusia is the same as on subcladogram 12 of the 648-step cladograms (Fig. 19) but that 
the Autoplusiina and the Plusiina have exchanged positions. 

However, the 647-step cladograms (which, given that the branch-swapping procedure ran to comple- 
tion, are most likely the shortest for this data set) are based upon the same characters (with one exception) 
as the previously rejected 648-step subcladogram 12. Both topologies unite the Argyrogrammatini, 
Plusiina and Autoplusiina using characters 83:0 — -> 3 and 1 16, and treat Rachiplusia as the sister-group of 
the Argyrogrammatini based on characters 14:0 — > 2 and 110. In addition, the latter relationship is 
supported on the 647-step cladograms by reversals in characters 18 and 148, both of which are highly 
homoplasious. Finally, the Plusiina are placed as sister-group to the Argyrogrammatini + Rachiplusia 
solely on the basis of character 162. This character (presence/absence of a subapical cornutus) is also highly 
homoplasious and was rejected as reliable in connection with the discussion of the relative positions of 
Agrapha and Plusiopalpa. Thus, the 647-step minimum-length cladograms can no more be accepted than 
those 648-step cladograms that include subcladogram 12. Therefore, they are rejected in favour of a longer 
cladogram but one in which the supporting characters are more reliable. 

Evaluation of the preferred cladogram 

Following from the decisions reached in the previous two sections, the preferred cladogram (Figs 27, 28) is 
648 steps in length and consists of subcladograms 2, 3, 5, 10 and 11; the outgroup topology that places the 
Stictopterinae closest to the Plusiinae; and the sections held in common by all the alternative 648-step 
cladograms. The first part of the evaluation of the (preferred) cladogram is character-oriented insofar as 
the emphasis is placed primarily upon the branches along which a given character changes. Each character 
will be taken in turn and its character state distribution on the cladogram discussed, with particular 







122 



IAN J. KITCHING 




Fig. 27 The preferred 648-step cladogram showing the branching sequence of the outgroups and the 
Plusiinae except for the Plusiini (see Fig. 28). The numbers on the nodes are the clade reference numbers 
used to identify clades in the evaluation of characters and taxa. 



SYSTEMATICS OF THE PLUSIINAE 



123 




Fig. 28 The preferred 648-step cladogram depicting the branching sequence of the genera of the tribe 
Plusiini. The numbers are the clade reference numbers. 



124 IAN J. KITCHING 

reference to any homoplasy, alternative optimizations and possible errors in coding and scoring states. The 
character consistency index (c) of each character is given in parentheses after the character number. The 
second part of the evaluation is taxon-oriented and examines each of the clades and its supporting 
characters in turn. The aim here is to determine to what degree each clade may be expected to be 
corroborated in the light of further data. Clades without formal names are referred to by numbers that are 
given on the cladograms in Figs 27, 28. 

Character analysis 

Wing pattern 

1 (0-333) The presence of raised scales in the forewing cell is one of the classic Hampsonian noctuid 
characters. Hampson (1913) noted their occurrence in the Plusiinae in Abrostola and Mouralia, and 
also in Chrysanympha, the sole species of which he placed in Abrostola. Raised scales in Chrysany- 
mphawere not observed by Eichlin & Cunningham (1978), nor were they in this study. This character 
state also occurs elsewhere in the Noctuidae, most notably as a defining character for the Sarrothripi- 
nae and as a secondary character for distinguishing the Stictopterinae from the Euteliinae. However, 
in this study, the forewing of Lophoptera (Stictopterinae) was found to be smooth, while raised scales 
were observed in Anuga (Euteliinae). As a result, 1 has a low consistency, being convergently 
derived in the Abrostolini, Stictoptera and Anuga. It weakly supports the monophyly of the former 
but more extensive sampling is required to adequately document its occurrence in the Stictopterinae 
and Euteliinae. 

2 (1-000) This character is apomorphic for the Abrostolini. Superficially similar patterns occur 
elsewhere (e.g. Paectes abrostoloides (Guenee)) but lack this element. 

3 (0.143) The Y-mark is one of the characteristic elements of the forewing pattern of the Plusiinae. 
Within the subfamily, it appears to be synapomorphic for the Argyrogrammatini + Plusiini, but with 
multiple reversals. In some instances {Anadevidia, Allagrapha), the lack of a Y-mark appears to be 
part of a general simplification of the wing pattern, while in others (Thysanoplusia, clade 22), it is due 
to further modification and elaboration of the pattern and possible overlaying of the Y-mark by more 
derived elements. The latter hypothesis is supported in Thysanoplusia by the observation that only T. 
intermixta and a few, apparently closely related, species lack the Y-mark (see character 9), while the 
remaining species (e.g. T. indicator (Walker), T. daubei) possess a more typical pattern including a 
Y-mark, albeit somewhat modified. Diachrysia (q.v.) merits further attention because in some 
species (e.g. D. leonina (Oberthiir)), the pattern is as simple as in Allagrapha, but in others (e.g. D. 
chryson, and more so, D. chrysitis and D. zosimi (Hiibner)), this simple pattern is further elaborated 
with large areas of golden-green metallic scales (see character 9). Within clade 22, Lamprotes 
appears to have regained the Y-mark. This result suggests that the small golden C-mark in this genus 
may not be homologous with the Y-mark. 

An alternative optimization of character 3, in which the Y-mark is convergently gained in clade 6 
(Argyrogrammatini less Anadevidia) and the Plusiini, with the lack in Anadevidia being treated as 
plesiomorphic, is rejected as a less satisfactory interpretation. 

4 (0.667) State 1 for this character is synapomorphic for the Argyrogrammatini and Plusiini, although 
there are independent reversals in Zonoplusia and Eutheiaplusia. State 4, in which the pink line is 
replaced by a white line, is independently derived from state 1 and is synapomorphic for Euchalcia 
and Adeva. There are alternative optimizations for states 2 and 3, the best hypothesising the former 
state to be a synapomorphy of Ctenoplusia and Acanthoplusia, with the latter condition being 
autapomorphic for Ctenoplusia. However, state 3 is not universal within Ctenoplusia, but may help 
delimit a subset of this genus including not only C. limbirena, but perhaps also such species as C. 
melanocephala (Moschler), C. micans (Dufay) and C. polycampta Dufay. 

5 (0-200) Scattered, individual black scales on the forewing appear to be convergently derived in all 
the genera that possess them. This was not unexpected for Erythroplasia, in which such scales are 
present in one of the species (E. rutulifrons) but absent in the other (E. pyropia). In addition, this 
genus is only distantly related to the other genera that possess these scales, most of which are 
members of the Euchalciina. Within the Euchalciina, state 1 has been independently gained in 
Polychrysia, Chrysanympha, Adeva and Pseudeva, suggesting that the correct homologies have not 
yet been discovered. Further sampling, however, is unlikely to improve matters. A cursory 
examination of the other species in the genera concerned showed that black scales were absent 
entirely in all species of Panchrysia, Euchalcia and Plusidia (providing Plusidia imperatrix Draudt is 
placed in Polychrysia, see below), and present in all Polychrysia and Pseudeva, although only weakly 
developed in P. palligera (Grote). Scattered black forewing scales can also be seen in the unanalysed 
Syngrapha parilis (Fig. 96). 






SYSTEMATICS OF THE PLUSIINAE 125 

6 (0-100) The presence or absence of the tornal scale tooth is a highly variable and homoplasious 
character. Although characteristic of the Argyrogrammatini and Plusiini, it is multiply lost and 
regained, with numerous alternative sequences of change. One such sequence in clade 4 postulates 
acquisition followed by subsequent loss in Agrapha, Anadevidia, Erythroplusia, Loboplusia, clade 
24 and clade 40, with regain in Palaeographa, Autographa bractea and Lamprotes + Pseudeva. The 
scale tooth is evidently a highly unreliable character. 

7 (0-500) Yellow hindwings occur scattered throughout the Plusiinae and seem to be partially 
correlated with an arctic-alpine habitat. In the taxa under study, they have been independently 
developed in Omorphina and Syngrapha + Caloplusia. Whilst yellow hindwings are characteristic of 
the group of species related to 5. devergens and C. hochenwarthi(e.g. C. tibetana Staudinger, S. ignea 
(Grote), S. microgamma (Hiibner), S. montana (Packard), S. parilis [although this last species has 
unique white hindwings]), they also appear to have been independently derived (Eichlin & 
Cunningham, 1978) in another group of syngraphas, which includes S. diasema (Boisduval), S. borea 
(Aurivillius), S. diversigna Ottolengui, S. orophila, S. sackenii (Grote) and 5. snowii (Edwards). 
Additional plusiines with yellow hindwings include Trichoplusia exquisita (Felder) (Fig. 45), 
Panchrysia dives (Eversmann), Rachiplusia virgula (Blanchard) and Autographa sunsoni Dod. This 
colour pattern may prove to be of use within Syngrapha (s.l.) in a wider study but should be treated 
with caution. 

8 (1-000) A hyaline hindwing base is apomorphic for the Stictopterinae. 

9 (0-333) Extensive patches of green-gold scales have been independently acquired by Chalcopasta, 
Thysanoplusia and Diachrysia. The former supports Hogue's (1963) contention that this colour 
pattern is convergent in the stiriines and plusiines. This analysis also demonstrates that the pattern 
similarity between Thysanoplusia and Diachrysia is also superficial. 

10 (0-500) Similarly, a less pronounced, general metallic sheen on the forewings is also convergent 
between Chalcopasta and the plusiines. 

1 1 (0-333) The black spot at the apex of forewing vein Mt, is interpreted as convergent in Stigmoplusia, 
Plusiotricha and Pseudoplusia + Chrysodeixis. 

12 (0-100) This character, the degree of silver scaling around the reniform stigma, has already been 
dismissed above, when choosing between subcladograms 5-8, as virtually useless for providing 
evidence of relationship. 

13 (0-500) The loss of the hair-like scales on the forewing cell underside is interpreted as convergent in 
Chalcopasta and the euteliines + stictopterines. 

Wing structure 

14 (0-600) Three independent transformation series appear to be present in this character, all of which 
treat the widespread state as plesiomorphic and all of which have alternative sequences of state 
change. The most parsimonious for each in terms of logical transformations is as follows. Within the 
Plusiinae, the Argyrogrammatini are characterized by the change — * 1 , while clade 6 is defined by 
change 1 — > 2. The change 0— > 2 has been convergently derived in Rachiplusia and Autoplusia. The 
Stictopterinae + Euteliinae are united by the transformation — > 3. There are then two further 
independent changes: 3 — > 4 characterizes Paectes, while 3 — > 5 is apomorphic for the Stictopterinae. 
The node uniting Brachionycha and Diloba can take a value of 0, 2 or 6. An independent origin of 
state 2 in Brachionycha and of state 6 in Diloba is supported by the presence of state in 
Brachionycha nubeculosa (Esper), Psaphida rcsumens Walker and Copivaleria grotei (Morrison). 

15 (0-500) As was expected, the transformation series of reduction in hindwing vein M 2 (the trifine/ 
quadrifine dichotomy) did not emerge non-homoplasious. The plesiomorphic state remains uncer- 
tain but the Plusiinae can be delimited by state 1 (although this is convergently derived in Diloba, and 
Mouralia possesses state 2); the Stictopterinae + Euteliinae by state 0; and Calophasia by state 3. 
Whatever the eventual outcome, it is certain that the trifine condition will be found to have been 
attained on several, if not numerous, occasions. 

16 (0-250) The apically-produced forewing at vein R 4 is interpreted as convergently derived in 
Lamprotes + Pseudeva, Panchrysia, Allagrapha and Diachrysia. 

17 (0-500) This venational character unites the Stictopterinae and Euteliinae but is reversed in Paectes. 

18 (0-200) The origin of vein M 2 in the hindwing is partially correlated with character 15 and the 
development of the trifine condition. It is even more highly homoplasious, with numerous converg- 
ences and reversals, which include independent derivations of state 3 from both state 1 and state 2. 
Many genera appeared intermediate between states 1 and 2 and this character is perhaps best 
disregarded. 

19 (0-200) The apomorphic state for the outline shape of the hindwing has been developed five times 



126 IAN J. KITCHING 

independently in Mouralia, Panchrysia, Magusa, Calophasia + Cucullia and Lamprotes + Pseudeva, 
although the precise condition in Panchrysia and Lamprotes is unclear. Much of the homoplasy may 
be due to its partial correlation with character 15. 

20 (0-667) The plesiomorphic state for the scale arrangement appears likely to be state 1, with state 2 
being a good synapomorphy for the Stictopterinae + Euteliinae + Magusa. State is interpreted as 
convergent in Omorphina and Brachionycha + Diloba. 

21 (1-000) The presence of a short distal section of forewing vein CuP is synapomorphic for Bra- 
chionycha and Diloba. 

Head 

22 (0-333) Lashed eyes characterize the Plusiinae and clade 51 of the outgroups, with reversals in 
Brachionycha and Chalcopasta. Very long lashes have been independently acquired in Abrostola, 
Cucullia and Diloba. 

23 (1.000) Bulbous eyes unite the Stictopterinae, Euteliinae and Magusa. 

24 (0-500) The prominent rim around the eyes has been convergently gained in Omorphina and 
Syngrapha + Caloplusia. This character is an alternative statement of the 'classic' character 'eye 
small and elliptical', and has been functionally correlated with a diurnal habit (Hardwick, 1970: 15; 
see also Kitching, 1984: 184). 

25 (T000) A lower frons that is scaled only at the edge unites the Stictopterinae and Euteliinae. 

26 (T000) Like character 23, character 26, the degree of prominence of the frons, unites the Stictopter- 
inae, Euteliinae and Magusa, and may only be an alternative formulation of that character. 

27 (0-667) Triangular pilifers are apomorphic for the Plusiinae, although they are convergently derived 
in Calophasia. Broad, reduced pilifers are autapomorphic for Diloba. 

28 (1-000) A dorsally delimited frons unites the plusiine tribes Abrostolini, Argyrogrammatini and 
Plusiini. 

29 (1-000) A convex occiput is apomorphic for the Plusiinae. 

30 (0-500) A narrow frons in the male is interpreted as convergent in Stictoptera and Anuga. 

Proboscis 

31 (1-000) A reduction in the number of apical sensilla on the proboscis characterizes the Plusiinae, 
while an independent total loss unites Brachionycha and Diloba. This latter condition is probably 
a direct correlate of the extreme reduction of the proboscis found in these two genera (character 
32). 

32 (0-667) As referred to above, the proboscis in Brachionycha and Diloba is very reduced and 
non-functional. Most outgroup genera have a somewhat short, stout proboscis but in one pair, 
Calophasia + Cucullia, it is longer and thinner. This condition is convergent upon that found in the 
Plusiinae. 

33 (0-667) Most outgroup genera have ridged styloconic sensilla apically on the proboscis. In the 
plusiines, these sensilla are smooth, as they are in Cucullia, by convergent derivation. The euteliines 
are characterized by the further elaboration of three of the ridges into 'wings'. Close examination of 
the sensilla of Stictoptera (Fig. 113) reveals partially-developed 'wings', thus strengthening the 
hypothesised sister-group relationship between the Stictopterinae and the Euteliinae. 

34 (1-000) State 1 for this character unites Magusa, the Stictopterinae and the Euteliinae. State 2 is 
found only in Brachionycha and Diloba and is functionally correlated with character states 31:0 and 
32:2; the lack of apical styloconic sensilla allowing setae to extend to the apex of the reduced 
proboscis. 

35 (1-000) Only in the Plusiinae are the strengthening bars of the proboscis semicircular to the apex. 
The apical nodules in the euteliines are restricted to the extreme tip of the proboscis and are preceded 
by very strong semicircular bars. However, these bars appear to be structurally different to those in 
the Plusiinae and are treated as non-homologous. 

36 (1-000) Dense, regular, basal microspines on the proboscis are characteristic of the euteliines. 

Labial palps 

37 (1-000) Smooth ventral edges to the labial palps unite the stictopterines, euteliines and Magusa. 

38 (0-500) Although the apomorphic state for this character defines the outgroup clade 51 (less 
Oncocnemis + Chalcopasta, in which it is reversed), the semi-erect scales in the four genera 
concerned are differently distributed and thus possibly not homologous. In Diloba and Cucullia, 
semi-erect scales are present on both the inner and outer surfaces; in Brachionycha, they are 
restricted to the inner surface; while in Calophasia, they occur only on the outer surface. 



SYSTEMATICS OF THE PLUSIINAE 127 

39 (0-500) A blade-like labial palp segment 3 is characteristic of the Euchalciina but is reversed in 
Euchalcia + Acieva. 

40 (0-500) A labial palp in which segment 1 is subequal in length to segment 2 is apomorphic for clade 51 
in the outgroups but is reversed in Calophasia. 

41 (0-250) An elongate labial palp segment 3 is interpreted as having been independently gained with- 
in the Euchalciina in Polychrysia, Chrysanympha and Lamprotes + Pseudeva. Plusiopalpa also 
has a superficially similar elongate segment 3 but in this genus, close examination reveals that 
the elongation is dorso-distal to the VRO (Fig. 157) and so not homologous with the general elon- 
gation found in the other four genera, where the VRO remains subapical in position (e.g. Fig. 
156). 

42 (0-250) An acutely pointed labial palp segment 3 has been independently acquired in Omorphina, 
Oncocnemis, Lamprotes + Pseudeva and Panchrysia + Chrysanympha. 

43 (0-500) There are two possible transformation sequences for this character on the cladogram. State 1 
can either be independently gained in the Stictopterinae and Paectes, or unite the Stictopterinae and 
Euteliinae, with subsequent reversal in Amiga. This latter optimization is favoured by the observa- 
tion that, while a third stictopterine, Nigramma, possesses state 1, a fourth, Odontodes, lacks it. 
Thus, in this slightly larger sample of genera, state 1 would have to be reversed in Odontodes (as this 
genus would form the sister-group of Lophoptera rather than be the basal stictopterine lineage; 
Holloway, 1985), and so it seems more likely that reversal has also occurred in Anuga. 

44 (0-500) A labial palp segment 3 that is subspherical appears to have been independently derived in 
Brachionycha and in Chalcopasta + Oncocnemis. 

Antennae 

45 (0-500) Very long ventral hairs on the antennae have been independently acquired in Stictoptera and 
Paectes. 

46 (0-667) Asymmetrically bipectinate antennae are characteristic of the Euteliinae, whereas symmet- 
rically bipectinate antennae have independently arisen in Brachionycha + Diloba and Lophoplusia. 

47 (1-000) The three states of this character are mutually exclusive to the three groups that have 
pectinate antennae: Lophoplusia (state 1), the Euteliinae (state 0), and Brachionycha + Diloba 
(state 2). The remaining genera were coded * and, as a result, the optimization procedure allowed 
every node in the cladogram between the three groups mentioned to take any value, 0, 1 or 2. Only a 
more detailed study of antennal morphology will determine whether any (or none) of these states is 
comparable to the condition found in those genera with simple antennae, and thus which, if any, is 
plesiomorphic. 

48 (1-000) Pectinations that arise basally on each segment may be synapomorphic for Brachionycha 
+ Diloba. Alternatively, pectinations that arise distally could be apomorphic for a large group 
of taxa, including the euteliines and plusiines, but are only expressed in a small minority of the 
included species. Only the discovery of pectinations in a taxon that normally has simple antennae 
might provide the required information to allow a decision to be made as to which state is 
plesiomorphic. 

External thoracic structure 

49 (0-500) A large thoracic crest is apomorphic for the Plusiinae but also occurs in Chalcopasta. 

50 (0-500) Vertically-held tegulae appear to have been convergently acquired by Polychrysia and 
Pseudeva. This was a difficult character to score, and partially-erect tegulae occur in other euchal- 
ciine genera (Lamprotes, Plusidia). More detailed examination may reveal that character 50 defines 
a much larger group. 

51 (1-000) A truncate dorsal tegular arm is apomorphic for the Abrostolini. 

52 (0-667) State 2 of this character displays a similar distribution to character 43, except in this instance, 
the apomorphic state can either unite the Stictopterinae and Euteliinae then reverse in Stictoptera, 
or be independently derived in the Euteliinae and Lophoptera. State 1 is autapomorphic for Omor- 
phina. 

53 (0-500) The colourless, microspinose extension to the ventral tegular arm is convergently derived in 
Stictoptera and Anuga. 

54 (1-000) Micospines along the inner edge of the ventral tegular arm are synapomorphic for 
Calophasia and Cucullia. 

55 (1-000) A ventral tegular arm that is produced on a lateral extension of the main tegular body is 
synapomorphic for Brachionycha and Diloba. 

56 (1-000) A broad marginal flange on the dorsal tegular arm may prove synapomorphic for the 



128 IAN J. KITCHING 

Plusiinae, Euteliinae, Stictopterinae and Magusa. However, only Brachionycha and Diloba truly 
lack a flange ; the members of clade 52 possess a very narrow flange , less than one scale-base wide , and 
thus further sampling may result in reinterpretation of this character. 

57 (1-000) The attachment point of the patagium is drawn out only in the Stictopterinae + Euteliinae. 

Legs 

58 (0-111) This character posed coding problems for those genera in which the foretibia was striped or 
mottled (Chrysanympha, Adeva, Cucullia, Diloba, Magusa, Stictoptera), or entirely white (Eosphor- 
opteryx). These taxa were coded *. This resulted in several alternative sequences of state change in 
the Euchalciina. Given that the foretibial white spot is frequently lost elsewhere in the subfamily 
(Zonoplusia, Allagrapha, Diachrysia, Plusia), the best optimization assigns state 1 to Adeva and 
independent losses to Phisidia and Euchalcia. Chrysanympha and Eosphoropteryx were optimized to 
bear state 1. In the outgroups, the white spot was convergently acquired by the euteliines and 
Chalcopasta. Cucullia, Diloba, Magusa and Stictoptera were treated as having state 0. 

59 (0-083) The colour of the dorsal surface of the hind tibiae and tarsi is the most homoplasious 
character in the data, with six gains and six losses, yet with only one ambiguous assignment of state to 
an internal node. The uniform white colour may have been independently gained by Omorphina and 
the Argyrogrammatini + Plusiini, or gained by the Plusiinae as a whole and lost in the Abrostolini. 
Convergent gains occur in Chalcopasta, Cucullia, Calophasia (following loss in clade 43) and 
Autoplusia (following loss in clade 34). The remaining reversals occur in Rachiplusia, Pseudoplusia 
and clade 9. Many genera proved difficult to code in that they had a generally unicolorous pattern in 
which the white apices to the segments were still faintly visible: Trichoplusia, Zonoplusia, Pseudo- 
plusia, Argyrogramma, Agrapha, Adeva and Syngrapha. They were originally coded 0. Had these 
taxa been coded 1 , the homoplasy would actually have been greater by one step. Character 59 is thus 
best dismissed as containing any reliable information. 

60 (0-500) The basitarsal comb on the male hindleg appears at first analysis to be a very labile character 
that is interpreted as having been multiply acquired and transformed within the Argyrogrammatini. 
State 1 is present in Dactyloplusia, Trichoplusia + Thysanoplusia and Ctenoplusia + Acanthoplusia, 
as an independent development in each. State 2 occurs as a transformation of state 1 in Ctenoplusia 
and as an independent derivation from state in Plusiotricha. This is not improbable because the 
comb in Plusiotricha consists of a single row of about 9 setae , while that in Ctenoplusia comprises only 
two setae (but see below). State 3 in Argyrogramma + Stigmoplusia also arises de novo. However, 
while two other species of Stigmoplusia (S. allocota Dufay and S. acalypta Dufay) also possess state 3, 
two other Argyrogramma (A. basigera and A. signata) lack combs entirely. Additionally, while all 
species of Agrapha lack a differentiated comb, in one species (an unnamed species in the BMNH 
collection; sp. A, see Agrapha, Clade analysis), the inner SV row of spines is basally longer and 
thinner. 

For the remaining genera, a more thorough survey was performed. All Acanthoplusia species 
examined (N = 4) agreed with A. tarassota. Most Ctenoplusia species (N = 30) had a short row of 2-7 
setae, agreeing with C. limbirena in setal form, if not exactly in number. C. aenofusa (Hampson) and 
C. phocea (Hampson) exhibited state 3 and this suggests that these two species may not belong in 
Ctenoplusia but may be more closely related to Argyrogramma and Stigmoplusia. C. epargyra 
(Dufay) and C. leucostigma (Dufay) lack both a tarsal comb and true blade-like setae on the valve 
(see character 133), and, likewise, perhaps should be excluded. The former owes its place in 
Ctenoplusia (Dufay, 1970a) to a superficial similarity to Ctenoplusia mapongua (Holland) and 
Trichoplusia sestertia (Fig. 44) (then in 'Plusia' s.l.), while the latter resembles C. phocea in habitus. 
Finally, C. albostriata (Bremer & Grey) and C. subchalybaea (Walker) (treated as a synonym of the 
previous species by Holloway, 1985), together with Trichoplusia oxy gramma and T. vittata, also lack 
a tarsal comb. The first three taxa, however, do bear modified setae along the ventral edge of the 
valve. Nevertheless, they may not belong in Ctenoplusia. If the above anomalous species of 
Ctenoplusia are excluded, then the genus becomes extremely well circumscribed by the form of the 
tarsal comb. The modified setae may then prove to be synapomorphic for the genera Ctenoplusia, 
Acanthoplusia, [Eutheiaplusia] and Plusiotricha (see character 133). 

Within Trichoplusia (N = 29, excluding T. oxy gramma and T. vittata) and Thysanoplusia (N = 9), 
eight species of the former (T. rectilinea (Wallengren), T. sestertia, T. exquisita, T. arachnoides, T. 
callista Dufay, T. photeina Dufay, T. dolera Dufay and T. gromieri Dufay) and three species of the 
latter (Thy. orichalcea, Thy.florina (Guenee) and Thy. viettei (Dufay)) lack tarsal combs. However, 
the vagueness of the limits of these genera renders a critical evaluation impossible until a species level 
cladistic analysis has been undertaken. 



SYSTEMATICS OF THE PLUSIINAE 129 

Overall, it should not be considered unlikely that tarsal combs will be shown to have evolved two or 
more times within the Argyrogrammatini, with multiple, independent subsequent losses. Neverthe- 
less, state 3 may prove to be synapomorphic for Stigmoplusia, (?part of) Argyrogramma and several 
possibly related species (ICtenoplusia aenofusa, C. phocea), while state 2 may be apomorphic for 
Ctenoplusia, or a large subset thereof. 

61 (0-600) State 1 for this character has been acquired independently three times, in the Stictopterinae, 
Plusiotricha and clade 17. The first of these is almost certainly convergent upon the plusiine condition 
as a result of logical dependence on the following character. Further suppression of the male 
foretibial spines (state 2) is synapomorphic for Argyrogramma and Stigmoplusia , while the most 
extreme modification is autapomorphic for Argyrogramma. 

62 (1-000) Reduced hind basitarsal spines are apomorphic for the Stictopterinae + Euteliinae, with 
total loss on the basal half of the segment occurring in the stictopterines. Spines are also suppressed 
on the other legs in species showing state 2 and this gives rise to a correlated coding for these genera 
for character 61. 

63 (1-000) Spining on the fore and midtibiae was treated separately from that on the hindtibiae (see 
character 64). Apical spines (state 1) were autapomorphic for Rachiplusia, while extensive spining 
(state 2) was autapomorphic for Calophisia. 

64 (0-429) Spines on the inner surface of the hindtibia are autapomorphic for Mouralia and form a 
separate transformation series from that concerning spines on the outer surface. Most instances of 
the latter appear to have been independently derived: state 1 in Autoplusia, Cornutiplusia and clade 
44; state 2 in Anagrapha and Rachiplusia. Only in Calophisia is state 2 derived from state 1 . However, 
as mentioned above, occasional spines may be observed in Plusia and Autographa. and so may prove 
apomorphic for the more extensive clade 38, with multiple subsequent suppression. Also, in 
Autoplusia, one leg of the male A. egena examined showed state 2, while the other leg, together with 
both legs of the female, displayed state 1 . Thus this character is also plastic in Autoplusia. Overall , the 
expression of character 64 is variable and should be treated with extreme caution when assessing 
degrees of relatedness. 

65 (0-750) The three forms of SD and D setae on the tarsi have been independently derived; state 1 in 
Diachrysia, state 2 in Polychrysia, and state 3, convergently, in Panchrysia and Lophoplusia. 

66 (T000) Hindlegs are shorter than midlegs only in the Stictopterinae + Euteliinae. 

67 (0-500) Ventral lobes on the claw are convergently absent in Oncocnemis and Diloba. 

68 (0-667) Elongate hindleg inner apical spurs occur in the Stictopterinae and Euteliinae. Two 
sequences of character state changes are possible. States 1 and 2 can be independently derived in 
Stictoptera and the euteliines respectively (the latter via state 1); or state 1 can unite the two 
subfamilies with a transformation to state 2 in the euteliines and a reversal to state in Lophoptera. 
Further sampling within these two subfamilies is required in order to decide between these two 
competing transformation series. 

69 (0-500) Foretibial claws have been independently derived in Brachionycha and Oncocnemis + 
Chalcopasta. 

Internal structure of the prothorax 

70 (1-000) A posteriorly recurved prothoracic furca characterizes the Euteliinae + Stictopterinae. 

71 (0-500) A ridge on the dorsal edge of the epimeron is convergent between the Euteliinae and 
Brachionycha + Diloba. 

Male abdomen A 1-7 

72 (0-500) The narrowly tapered abdomen occurs in Stictoptera and the euteliines. As for character 17, 
this distribution can be accounted for by a gain followed by a loss, or by two independent gains. 

73 (1-000) State 3 forms an independent transformation, characterizing the Stictopterinae + Eute- 
liinae. There is some ambiguity regarding the fitting of states 1 and 2. The most morphologically 
parsimonious sequence treats state 1 as apomorphic for the Plusiinae less the Omorphinini, in which 
the ventro-lateral edges of the internal flange on A2 are expanded internally. This condition is 
retained in the Abrostolini , but in the higher Plusiinae (clade 3) , the lobes of the flange are even more 
highly developed, meeting medially but with a ventral notch. 

74 (1-000) Internal longitudinal flanges on St2 are autapomorphic for the Euteliinae. Holloway (1985) 
stated that these structures may be correlated with the resting position of the adults, in which the 
wings are longitudinally folded and the abdomen is recurved dorsally. 

75 (1-000) Components of the trifine hair pencils are found, in this study, only in Cucullia and Magusa, 
where they are interpreted as independently derived. Far more taxa need to be examined before any 



130 IAN J. KITCHING 

meaningful information can be extracted from this frequently highly labile (Birch, 1972b) character 
complex. 

76 (1-000) Postero-lateral sclerotizations on St7 are apomorphic for clade 6 - the Argyrogrammatini 
less Anadevidia. They are only just discernible in Pseudoplusia and Eutheiaplusia, a condition that 
may be correlated with the reduced hair pencils and subdorsal tufts of these genera (see below). 

77 (0-429) The presence of dorso-lateral hair tufts on segments A5-7 characterizes clade 6, with 
independent losses in Zonoplusia and Pseudoplusia. Additional tufts on segments A(2-)4 are 
synapomorphic for the members of clade 12, but with subsequent loss in Acanthoplusia + Ctenoplu- 
sia. The presence of distinct additional tufts on A4 is autapomorphic for Chrysodeixis. As alluded to 
above, it may eventually be possible to subdivide state 2 and extract more information, whence 
further resolution within the Argyrogrammatini may be forthcoming. 

78 (1-000) As was originally suspected, the three forms of enclosed hair tufts on A7 are independently 
derived in Dactyloplusia (state 1), Ctenoplusia (2) and Argyrogramma (3). 

79 (T000) This character (distinct edges to the tufts on A5 and A6) has been discussed earlier in 
connection with the decision between subcladograms 1 and 2. 

Male St8 and ventral hair pencils 

80 (0-600) This character is now shown to consist of at least two independently derived forms of hair 
pencil. State 3 characterizes the Stictopterinae + Euteliinae + Magusa, but is also found in 
Calophasia + Cucullia. The hair pencils coded by states 1 and 2 are restricted to the Plusiinae. The 
former unites the members of clade 2 (the Plusiinae less Omorphina), while state 2 has been 
independently derived in Dactyloplusia and the argyrogrammatine clade 10. There is some evidence 
to suggest that the conditions of St8 found in Zonoplusia, Pseudoplusia and Chrysodeixis may 
eventually be recognized as reduced forms of state 2, whence this feature would become apomorphic 
for clade 6. Several other taxa also show partial reductions of the hair pencils and St8. 

81 (1-000) The plesiomorphic state for the stictopterine type of hair pencil is interpreted as state 3. This 
causes some problems in the optimization of states on the internal nodes of the outgroups because 
Brachionycha, Diloba, Chalcopasta and Oncocnemis all lack any type of hair pencil and were coded 
*. However, given the ambiguity in the outgroup relationships and their relatively distant affinities, 
this is not a serious uncertainty in the current study. Within clade 47, state is probably plesiomor- 
phic, with states 2 and 1 being autapomorphic for Magusa and Anuga respectively. Given that 
Magusa may not represent the immediate sister-group of the Stictopterinae + Euteliinae (see clade 
47, Clade analysis), this sequence of state transformation seems more prudent than the alterna- 
tives. 

82 (1-000) Similarly, character 82 does not apply to clades 53 and 55. Thus state is apomorphic for 
Calophasia + Cucullia, and state 1 for clade 47. State 2 is autapomorphic for Stictoptera. Anuga, 
coded * , is predicted to have state 0. 

83 (0-313) This character, the degree of development of the male sternal hair pencils, proved, as 
expected, to be highly homoplasious, with numerous alternative character state optimizations. The 
sequence chosen assumed that state 0, as seen in Abrostola, is plesiomorphic for the Plusiinae (less 
Omorphina). The large single pocket (state 5) of Mouralia is autapomorphic within the Abrostolini. 
The initial transformation -h> 3 (well developed, separate pockets) characterizes the Argyrogram- 
matini + Plusiini. Within the Plusiini, the pockets are reduced to state again in the Euchalciina. This 
confirms that state in this subtribe is the result of secondary loss and is not due to plesiomorphic 
absence, as in Abrostola, providing that hair pencils have not been independently gained in the 
Argyrogrammatini and Autoplusiina + Plusiina. A change to state 2 (partially fused pockets) 
characterizes clade 30 (the Autoplusiina less Rachiplusia) but with a reversal to state 3 in Autoplusia 
itself. The Plusiina are characterized by the change 3 — > 1 (reduction of pockets to slight invagina- 
tions), with independent reversals to states 3 and in Plusia and Syngrapha respectively. 

The situation is less simple in the Argyrogrammatini. A parallel change to state 2 (as in clade 30) 
occurs to unite the members of clade 6. Within clade 8, there are independent transformations to 
states and 5 in Pseudoplusia and Chrysodeixis respectively. The form of the hair pencil is then even 
more highly modified in the more derived members of the latter genus, as outlined above (Terminol- 
ogy). State 4 (fused pockets) is apomorphic for clade 17, with a reversal to state 3 in Stigmoplusia. 
There are also parallel changes from state 2 to state 3 in Acanthoplusia and Trichoplusia + 
Thysanoplusia. 

The sternal hair pencils are obviously highly variable and maybe labile structures. However, the 
distribution of states and homoplasy is very far from random and more extensive sampling could 
result in this complex proving very useful. 



SYSTEMATICS OF THE PLUSIINAE 131 

84 (0-250) A sclerotized bar on the hair pencil pockets is apomorphic for clade 10 but with subsequent 
independent losses in Plusiopalpa, Acanthoplusia and Thysanoplusia. 

85 (1-000) Differentiated hair pencil insertions are synapomorphic for Argyrogramma and Stigmoplu- 
sia. 

86 (0-333) As discussed above, accessory pockets on the preferred cladogram must be developed in 
clade 14 but then independently lost in Plusiopalpa and Ctenoplusia. 

87 (0-500) Short, outer scales on the hair pencils also characterize clade 14, but with a reversal in clade 
17. 

88 (0-500) Long inner tufts of hair pencil scales are convergently derived in Agrapha and Argyro- 
gramma. 

89 ( 1-000) Secondary hair brushes are synapomorphic for the members of the Argyrogrammatini . 

90 (1-000) An anterior flange on the lateral arms of St8 is restricted to Argyrogramma and Stigmo- 
plusia. 

91 (0-500) Lateral arms of St8 that twist dorsally over the hair pencils are apomorphic for the 
Argyrogrammatini less Anadevidia, but the change is reversed in Pseudoplusia, which also has much 
reduced hair pencils. Further, the development of a deep basal concavity characterizes clade 12 but 
with subsequent reversal in Plusiopalpa. 

92 (0-750) This character was also discussed in detail above, with regard to subcladograms 1 and 2. 
State 1 is convergently derived in clade 12 and Dactyloplusia; state 3 is apomorphic for clade 14 and 
this transforms to state 2 to unite Argyrogramma and Stigmoplusia. 

93 (0-600) The form of fusion of the apices of the dorsal arms of St8 is fairly homoplasious. State 2 (in 
which the arms fuse into a rounded apex) is apomorphic for clade 10, with further transformations to 
state 3 (transversely oriented dorsal arms), uniting Argyrogramma and Stigmoplusia, and state 1 
(converging dorsal arms), autapomorphic for Thysanoplusia. However, state 1 occurs outside clade 
10 in clade 7, with a reversal to state in Pseudoplusia. This character has probably not been coded 
correctly and further sampling of taxa may clarify matters. The form of St8 will almost certainly prove 
to be highly important in the future critical resolution of the interrelationships of the taxa within the 
Argyrogrammatini . 

94 (0-750) There are several alternative optimizations for the internal node states for this character too. 
The most probable, on present evidence, interprets state 2 (presence of broad, flat, 'greasy'-looking 
scales ventrally between the dorsal arms of St8) as apomorphic for clade 6. There are subsequent 
transformations to state 3 (scales fused into a flat plate) in Argyrogramma + Stigmoplusia, to state 
(a reversal) in Pseudoplusia, and to state 1 (broad, erect, spatulate scales) in Chrysodeixis (which is 
further modified in the more derived species of this genus; see above). Superficially similar scales to 
those in Chrysodeixis occur in Anadevidia, but in the latter genus, they were interpreted as the scales 
of the secondary hair brush. Thus they are convergent upon the condition found in Chrysodeixis, 
which bears normal secondary hair brushes in addition to the apical spatulate scales. 

95 (0-500) Long, fine scales ventrally between the lateral arms of St8 characterize clade 30, although 
they are subsequently lost in Sclerogenia. 

Male genitalia - uncus, tegumen & anal tube 

96 (0-500) A ventrally flattened uncus is considered to have arisen independently in Paectes, Calopha- 
sia and Brachionycha + Diloba. State 2, a subspherical uncus, is autapomorphic for Anuga. 

97 (0-250) A straight uncus has been derived independently four times: Chalcopasta, Chrysodeixis, 
Stigmoplusia and clade 44. 

98 (0-500) The uncus has become distally dilated dorsally independently in Agrapha and Argyro- 
gramma. 

99 (1-000) A notch between the tegumen and uncus is characteristic of the Plusiinae less the Omorphi- 
nini. A further production of the tegumen apex unites Allagrapha, Macdunnoughia and Puriplusia. 

100 (1-000) A concave base to the uncus is an apomorphy of the Abrostolini. 

101 (1-000) A tegumen with a distinct angle is synapomorphic for the members of the Euchalciina. The 
apomorphic state is weak in Panchrysia and Pseudeva, and present in only one of the two Euchalcia 
specimens examined. 

102 (1-000) An uncus that is medially flattened and expanded is apomorphic for the Abrostolini. 

103 (1-000) The presence of a distinct median suture on the uncus is also apomorphic for the Abrosto- 
lini. 

104 (0-500) Widely divergent tegumenal arms are convergently derived in Antoculeora and Erythro- 
plusia. 

105 (0-111) The origin of the anal tube with respect to the LASTs and uncus base is highly homoplasious. 



132 IAN J. KITCHING 

This character seems generally to be correlated with the overall dorsal appearance of the tegumen 
apex, LASTs and uncus base. There is a general trend, which is difficult to quantify, that as the anal 
tube moves anteriorly, the apex of the tegumen becomes elongate and the outline of the tegumen — » 
LASTs — > uncus smoothes out. Until such time as the morphology of this complex region is clarified, 
the best optimization for the development of the apomorphic state may be to postulate independent 
gain wherever possible. Thus, state 1 may have arisen separately in Chalcopasta, clade 38 (with 
reversal in Plusia), Anadevidia, Agrapha, Argyrogramma, Trichoplusia and clade 7. In the last of 
these, the anal tube of Chrysodeixis chalcites arises at the uncus base but this condition appears to be 
the end product of a reversal series. The 'plesiomorphic' species, C. argentifera, has an anterior 
origin of the anal tube (state 1), as in Dactyloplusia and Pseudoplusia, while the 'intermediate' C. 
illuminata shows an intermediate position. The 'derived' C. kebea (and C. chalcites), as defined by an 
arrowhead-shaped St8 (see Terminology), have reverted totally to the plesiomorphic condition for 
character 105. 

106 (0-500) The fusion of the LASTs to the underside of the tegumen has occurred independently in the 
Argyrogrammatini + Plusiini and in Chalcopasta. 

107 (0-200) Subdorsal bars on the anal tube have been gained on several occasions: Chrysanympha, 
Lophoplusia, the Abrostolini, and clade 47 (with subsequent loss in the euteliines). Within the latter, 
an alternative optimization hypothesises independent gain in Magusa and the stictopterines. 
However, extreme modification of the LASTs in Paectes and their complete absence in Anuga 
suggests that loss of the bars in the euteliines is a more satisfactory hypothesis. 

Male genitalia - vinculum & pleurite 

108 (0-125) The presence of a ventral tooth on the vinculum is also a highly homoplasious character 
state. It has been acquired by clade 39 (with no reversals). Erythroplusia, Sclerogenia and the 
Argyrogrammatini. Within the latter, the tooth has been lost four times, in clades 7 and 13, and in 
Acanthoplusia and Argyrogramma. The tooth represents the insertion point of the S2 muscle (Bayer, 
1965), which, together with other muscles, retracts the genitalia into A8. Bayer noted the absence of 
S2 in Chrysodeixis acuta, a species that also lacks a tooth. Thus, an extensive investigation of genital 
musculature, and its correlation with the presence/absence of the vincular tooth, might help clarify 
the homoplasy exhibited by character 108. (For an alternative musculature nomenclature, see 
Tikhomirov, 1979.) 

109 (0-333) A vinculum that is S-shaped in lateral view has been independently gained in clades 21 
(weakly expressed) and 39 and in Cucullia. Since the SI muscle inserts dorso-apically on the vinculum 
(Bayer, 1965), this shape could also be functionally correlated with the musculature for retracting the 
genitalia. 

110 (0-200) A Y-shaped vinculum has been convergently derived in Paectes, Rachiplusia and clade 6 
(with subsequent reversal in Dactyloplusia and Trichoplusia). This character is relatively trivial and 
probably of low importance. 

111 (0-500) An attenuated spine-like vinculum is apomorphic for Chrysodeixis + Pseudoplusia. Stigmo- 
plusia has a produced vinculum but with a rounded apex, a condition shown here to be non- 
homologous. 

112 (0-750) Dorso-lateral emarginations (state 1) of the vinculum arms have been independently 
acquired by Thysanoplusia and clade 14. Within the latter, the emarginations are expanded into 
sharp points (state 2) in Stigmoplusia. The blunt hooks (state 3) of Plusiotricha represent an 
independent, non-homologous derivation from state 0. 

113 (0-333) A broad, flat, cone-like vinculum is regarded as apomorphic for the members of clade 31 but 
is reversed in the included clade 35. It is also independently derived in Anagrapha (but see Clade 
analysis). 

114 (0-500) A flattened, entire, conical vinculum is convergently derived in Rachiplusia and the 
stictopterines + euteliines. Originally scored *, Stictoptera was optimized to possess state 1. This 
suggests that further work in the Stictopterinae should show the condition in this genus to be 
derivable from state 1. 

115 (1-000) Distally convergent vinculum arms are apomorphic for the Euteliinae. 

116 (0-333) A distinct, sharp, basal notch on the pleurite is apomorphic for the Autoplusiina + Plusiina, 
although it is convergently present in Anadevidia and reversed in Antoculeora. The latter condition 
was commented upon above, when discussing the position of Rachiplusia and the branching 
sequence of the subtribes of the Plusiini, and is interpreted as being correlated with the highly 
modified male genitalia found in Antoculeora (see also characters 117 and 118). 

117 (0-500) A broad, flat lateral flange on the pleurite is considered to be an apomorphy of the Plusiini. 



SYSTEMATICS OF THE PLUSIINAE 133 

Like character 116, the apomorphic state is lost in Antoculeora. According to Bayer (1965), this 
flange represents the insertion point for the genitalia retraction muscle S3. 

118 (0-667) State 1, in which the pleurite lies along the antero-dorsal edge of the tegumen, is interpreted 
as plesiomorphic. State 2, where only the apex of the pleurite is in contact with the tegumen, is 
autapomorphic for Antoculeora, while Magusa and Calophasia appear to have independently 
acquired state 0, in which the apex of the pleurite articulates on the inner edge of the tegumen in a 
pocket. However, * codings for Diloba, the stictopterines and the euteliines render this conclusion 
open to question. 

119 (0-500) Partial fusion of the pleurite and VAVB has convergently arisen in Plusiotricha and 
Ctenoplusia + Acanthoplusia (but see characters 60 and 133, and Clade analysis). 

120 (1-000) A very narrow pleurite closely appressed to the antero-dorsal edge of the tegumen is 
synapomorphic for Chrysodeixis and Pseudoplusia. 

121 (0-500) The three states of this character are treated as independent derivations, all of which occur 
twice. A very short or missing VAVB (1) characterizes the Plusiini and Stictopterinae. A VAVB 
modified into a blunt lobe (2) arose convergently in Abrostola and Argyrogramma. In Calophasia 
and Oncocnemis, the VAVB is missing and the tegumen base is produced below the pleurite to 
approach the vinculum apex (3). 

122 (1-000) Socii are apomorphic for the euteliines. 

123 (0-500) Complete fusion of tegumen and vinculum arms has, as suspected a priori, occurred in 
independently in Diloba and the Stictopterinae + Euteliinae. 

Jaxta & anellus 

124 (1-000) The apex of the juxta is fused to the aedeagus ventrally only in the Abrostolini. 

125 (1-000) Apical thickening of the juxta is also apomorphic for the Abrostolini. 

126 (1-000) Fusion of the juxta, by a bar, to the inner edge of the sacculus near the base of the clavus 
occurs only in the Abrostolini. 

127 (0-429) A parallel-sided notch has developed apically on the juxta three times in the Plusiinae; in 
Ornorphina, Chrysodeixis and the Plusiini. Within the latter, it has been lost in Allagrapha and 
independently modified into a subapical process twice, in Anagrapha and clade 44. The further 
elaboration of this process into a sharp spine is synapomorphic for Syngrapha and Caloplusia. 
Although the character is homoplasious overall, state 3 may allow some resolution within Syngrapha 
(s.l.). The sharp spine is present only in a subset of those species with yellow hindwings (see character 
7) allied to S. devergens and Caloplusia hochenwarthi: S. ignea, S. microgamma, and S. alticola. The 
spine is absent in S. montana and represented by a 'strong point, but not a distinct, sclerotized spine' 
in S. parilis (^Eichlin & Cunningham, 1978). However, similar structures occur elsewhere; S. 
angulidens (Smith) has a sharp apical point, S. sackenii shows a similar structure to 5. parilis, and 
many other species exhibit a weak, blunt projection . Nevertheless, used with caution, the form of the 
juxta apex should prove useful in elucidating relationships within Syngrapha and maybe the Plusiina 
as a whole. 

128 (0-400) A scobinate juxta apex is apomorphic for clade 7, with distinct thorns autapomorphic for 
Pseudoplusia. State 1 has also been independently derived in Thysanoplusia and, as initially 
suspected, Plusiopalpa and Cucullia. 

129 (1-000) A weakly sclerotized, crescentic juxta is autapomorphic for the Euteliinae. 

130 (0-333) For this character, the outgroups were all coded *. The plesiomorphic state in the Plusiinae 
was interpreted to be state 1 , in which the juxta is fused to the saccular flanges. The juxta has become 
free three times independently, when saccular flanges are also present, in Agrapha, Pseudoplusia + 
Chrysodeixis and Argyrogramma + Stigmoplusia. Interestingly, all those plusiine taxa that lack 
saccular flanges (Abrostolini, Trichoplusia, Autoplusia) were optimized to display state 1 - juxta 
fused to flanges! This result suggests that a recoding of this character, incorporating the next (q.v.), 
will be necessary if any sense is to be made of the feature in future studies. 

Male genitalia - valve 

131 (0-250) The presence of saccular flanges characterizes the Plusiinae, with independent losses in the 
Abrostolini, Trichoplusia and Autoplusia. The implication for the previous character is that, had 
saccular flanges not been lost in these taxa, then they would have been fused to the juxta. 

132 (1-000) The presence of a rounded lobe between the valve bases, formed from the fused saccular 
flanges, is synapomorphic for Panchrysia and Chrysanympha. 

133 (0-500) Blade-like setae are interpreted here as convergently derived in Ctenoplusia + Acanthoplu- 
sia and Plusiotricha. As noted above (character 60), there appear to be problems regarding the 



134 IAN J. KITCHING 

homologies of this and similar setal modifications. In the present context, the setae coded by state 1 
are strongly sclerotized, regularly arranged in a single row along the ventral edge of the valve and 
shaped like short, broad, curved blades. In Ctenoplusia albostriata and Trichoplusia oxygramma 
(McDunnough, 1944), the ventral valval setae are weakly sclerotized and narrow, and thus may not 
be homologous with the setae found in typical ctenoplusias. Further work is necessary to decide 
whether this group of species truly belongs in Ctenoplusia or represents, say, the sister-group of 
Ctenoplusia (+ other genera?). Similarly, C. epargyra and C. leucostigma only have spiniform hairs 
on the ventral edge of the valve, which, in conjunction with their lack of a tarsal comb, argues for 
their exclusion from the genus. The setae in C. phocea are also weak and narrow. Additionally, they 
are in multiple rows basally, unlike other Ctenoplusia species. C. phocea also has a tarsal comb 
consisting of multiple rows of setae (60:3), an elongate vinculum (111:1) but with a rounded apex, 
and moderately strong dorso-lateral emarginations on the vinculum arms (112:1), strongly suggest- 
ing a relationship with Stigmoplusia. Finally, C. aenofusa is enigmatic. It has highly modified and 
autapomorphic setae along the ventral edge of the valve. They are strongly sclerotized and in a single 
row. However, the distal setae are narrow and strongly angled, while the basal setae are apically 
spatulate. Also, the setae on the basal half of the sacculus are borne on small marginal projections, 
unlike most ctenoplusias. As stated above, the tarsal comb is of the form found in Argyrogramma and 
Stigmoplusia, and C. aenofusa, too, may have its true affinities with these genera. 

All Acanthoplusia species possess blade-like setae. Dufay (19706) and Holloway (1985) state that 
these are missing in A. vermiculata Dufay, but recessed scale bases along the ventral edge of the 
valve, typical of Ctenoplusia and Acanthoplusia (and also Plusiotricha), are visible in the holotype. It 
must be assumed that the blade-like setae, which are often caducous in many species, have been 
accidentally removed before or when the specimen was prepared and mounted. 

Two other genera in clade 14 have modified valval setae. In Stigmoplusia, although the type 
species lacks such setae, three other species (S. allocota, S. acalypta, S. megista Dufay) all possess 
single, short, conical, somewhat flattened setae apically on the valve digitations. In Agrapha, the 
ventral edge of the valve is dorso-ventrally flattened and covered with a dense mat of short, upright, 
curved, flat scales. 

There is obviously a great deal of useful information in these valval setae within clade 12 but an 
exhaustive species-level analysis will be necessary to extract the maximum information. 

134 (1-000) A spatulate lobe arising from the sacculus is autapomorphic for the Abrostolini. 

135 (0-500) A uniseriate corona has been convergently derived in Magusa and clade 52. 

136 (0-500) The presence of basally-directed setose scales on the valve apex is synapomorphic for the 
Argyrogrammatini. Similar scales in Mouralia are interpreted as convergently derived. In Anadevi- 
dia peponis , the valval setae are extremely persistent and very numerous. However, basally-directed 
scales are present in A. hebetata (Butler) (Chou & Lu, 1978: fig. 2), in the position occupied by the 
caducous setae of the other argyrogrammatine genera, whereas no such setae occur anywhere in the 
Plusiini. 

137 (0-500) Setae arising from ventral marginal prominences on the valve are interpreted as apomorphic 
for clade 32, but with an independent derivation in Omorphina. 

138 (0-500) A spatulate apex (cucullus) to the valve is treated as synapomorphic for the members of 
clade 12 but with reversal in clade 17. The weak nature of the subapical constriction in Eutheiaplusia, 
together with weak narrowing in Trichoplusia, Thysanoplusia, Dactyloplusia and Pseudoplusia, 
suggests further sampling may reveal a very gradual transformation series leading from an uncon- 
stricted valve to the extreme condition seen in Ctenoplusia. 

139 (0-500) The clavus base is obscured by an overfold of the sacculus in Rachiplusia and clade 32 
convergently. 

140 (1-000) A broad, blunt point on the ventral edge of the valve unites Plusiotricha and Eutheiaplusia. 

141 (0-250) The saccular spine is convergently derived in Diloba and clade 30, but within the latter, it has 
been lost in Autoplusia and Loboplusia. The absence in Autoplusia egena is almost certainly a true 
loss (rather than plesiomorphic absence) since the spine is present in both A. olivacea and 
'Syngrapha gammoides. 

142 (0-333) A saccular sheet, in some form, is characteristic within the Plusiinae of clade 31, although it 
has subsequently been lost in Loboplusia. Further modification to form a hook unites Sclerogenia, 
Autoplusia and Antoculeora. Convergent occurrences of an apical saccular process appear in 
Lophoptera, Calophasia and Magusa. 

143 (0-500) The form of the clavus (characters 143-145) is highly homoplasious. Its presence on a 
distinct peninsula (state 3) is probably plesiomorphic for the Plusiinae. The peninsula is convergently 
lost in the Abrostolini and Anagrapha, so that the dorsal and ventral margins of the clavus are more 



SYSTEMATICS OF THE PLUSIINAE 135 

or less level (state 2). However, this state arose de novo from state in Cucullia. The clavus is further 
modified into a broad lobe in Abrostola, but this is convergently derived from an unmodified saccular 
margin in Chalcopasta + Oncocnemis. 

144 (0-133) This character, the degree of development of the clavus in the Plusiinae, was that used by 
McDunnough (1944) to subdivide the subfamily into three genus groups. The more extensive 
sampling of genera of the present study has shown this view to be far too simplistic. A fully 
developed, tubular clavus (state 0) is interpreted as plesiomorphic, because it is the form that occurs 
in the only outgroup with a distinct clavus {Cucullia). Multiple, independent reduction seems to be 
the rule, but some reversals (i.e. clavi becoming longer and better developed) do occur. Reduction to 
a short tubular clavus has occurred in the Argyrogrammatini, clades 24 and 43, Anagrapha, 
Loboplusia, Stigmoplusia (after a reversal in clade 15, see below) and Omorphina. Further 
reductions to a setose bump have taken place in clade 26, Lophoplusia, Plusiotricha and Agrapha. 
Reversal from state 2 to 1 occurs only in Lamprotes, or alternatively (and this, surprisingly, is the only 
internal node state ambiguity), state 2 could have arisen independently in Plusidia and Pseudeva. 
However, as Lamprotes exhibited a clavus form somewhat intermediate between states 1 and 2 (see 
Characters), this is not too problematical. Reversals from state 1 to occur only in the Argyrogram- 
matini but thrice, in Pseudoplusia + Chrysodeixis, Trichoplusia + Thysanoplusia and in clade 15. As 
seems to be true of all the widely quoted 'classic' key characters (see also, for example, characters 6, 
15 and 64), the form of the clavus is highly unreliable as an indicator of cladistic relationship. 

145 (0-333) An elongate, flattened peninsula has been convergently derived in Erythroplusia, Macdun- 
noughia + Puriplusia and Antoculeora. Interestingly, however, all four of these genera are members 
of the Autoplusiina. 

146 (1-000) A blade-like valve is apomorphic for the Plusiina. 

147 (0-500) More or less oval valves are apomorphic for clade 21 of the Euchalciina but also occur 
convergently in Omorphina. 

148 (0-200) The presence or absence of the harpes is also very homoplasious, with presence being 
interpreted as plesiomorphic. Five independent losses appear to have occurred, in the euteliines, 
Stictoptera, Rachiplusia, Anadevidia and clade 34. An alternative optimization in clade 48 hypothe- 
sises loss as uniting the Stictopterinae and Euteliinae, with a subsequent regain in Lophoptera. 

149 (0-250) It was suggested a priori that the various forms of inwardly-pointing hooked harpes would 
prove to be convergently derived and therefore probably non-homologous. This result was obtained, 
with independent derivations occurring in clade 44, Trichoplusia, Dactyloplusia and Agrapha. 

150 (1-000) Inwardly-curving hooked harpes are apomorphic for Macdunnoughia and Puriplusia, with 
further development of a basal, shorter, upcurving hook autapomorphic for the former genus. 
Interestingly, Macdunnoughia crassisigna (Warren) shows an intermediate condition in which the 
second hook is represented by a small sharp point. 

151 (1-000) Deeply cleft valves are autapomorphic for the Stictopterinae. 

152 (1-000) Trapezoid valves are autapomorphic for the Euteliinae. 

153 (1-000) Valves that, when closed, lie ventrally or subventrally are synapomorphic for the Euteliinae 
and Stictopterinae. 

154 (0-333) Transtillae that are fused and produced posteriorly to a broad, blunt point are convergently 
derived in Rachiplusia and clade 22 (with reversal in the included clade 26). 

155 (0-500) Bifurcate transtillae, discussed earlier in connection with the position of Loboplusia, are 
hypothesised to have arisen independently in Erythroplusia and Sclerogenia. 

Male genitalia - aedeagus 

156 (0-500) A thin, small aedeagus is apomorphic for clade 47 but reversed in Paectes. 

157 (0-667) A bulbous coecum (state 2) is synapomorphic for the Plusiini and Argyrogrammatini. In the 
latter tribe, the coecum can become very enlarged (e.g. 'Plusia' megaloba Hampson [Holloway, 
1985: 300]; Ctenoplusia aenofusa [Chou & Lu, 1978: fig. 4]), possibly in order to accommodate an 
extremely long vesica when retracted (Holloway, 1985) . A dorso-basal origin of the cuticular simplex 
appears to have been convergently acquired by Magusa and the euteliines. 

158 (0-400) Restriction of the aedeagus sclerotization to a ventral band is synapomorphic for the 
Argyrogrammatini and Plusiini, although there has been a reversal to a uniformly sclerotized 
aedeagus in Autoplusia + Antoculeora. Further reduction of the sclerotization to a narrow, 
well-defined ventral strip is apomorphic for the Argyrogrammatini but has been independently 
acquired by the autoplusiine genera Macdunnoughia and Erythroplusia. 

159 (0-500) A vesica that is entirely covered with small spines unites Plusiotricha and Eutheiaplusia, but 
also occurs in Thysanoplusia. 



136 IAN J. KITCHING 

160 (1-000) A strongly serpentine inverted vesica is also synapomorphic for Eutheiaplusia and Plusio- 
tricha. 

161 (1-000) The presence of a small oval sclerotized plate at the base of the ductus ejaculatoris is 
synapomorphic for the Stictopterinae and Euteliinae. 

162 (0-111) The distribution of states of this character was discussed in part above in connection with the 
positions of Agrapha and Plusiopalpa. The presence/absence of a subapical cornutus (SAC) is highly 
homoplasious, with several ambiguous assignments of character states to the internal nodes. 
Initially, the presence of a SAC is synapomorphic for the Argyrogrammatini + Plusiini. However, 
there have been numerous independent losses, in clades 7, 30 and 44, and in Lamprotes and 
Eosphoropteryx. Loss can also be interpreted as having occurred in clade 12, with subsequent regains 
in Agrapha and Argyrogramma + Stigmoplusia, but several alternative sequences of change are 
possible, including independent losses in Plusiotricha + Eutheiaplusia, Ctenoplusia + Acanthoplusia 
and Plusiopalpa. Given that Eutheiaplusia may have a SAC (see Characters) and many ctenoplusias 
definitely do (see 648-step cladograms: Agrapha and Plusiopalpa), multiple independent losses may 
represent the most likely events. 

163 (1-000) An apically swollen SAC is synapomorphic for Panchrysia and Chrysanympha. 

164 (1-000) A SAC arising from a circular concave plate is synapomorphic for Euchalcia and Adeva. 

165 (0-500) A SAC originating on a lateral diverticulum has arisen convergently in Polychrysia and 
Diachrysia. 

166 (1-000) This character (grooved/'winged' SAC) was discussed above in detail with regard to the 
branching sequence of Plusia, Cornutiplusia and Autographa. 

167 (T000) The presence of an invaginated sac in the vesica is autapomorphic for clade 6. 

168 (T000) A vesica in which the base is internally granular is autapomorphic for clade 24. 

169 (T000) Only the Abrostolini possess a ventral, apical, blunt point to the vesica sclerotization. This is 
autapomorphically modified into a hook in Abrostola. 

170 (1-000) A large membranous RHS diverticulum on the vesica, bearing cornuti is synapomorphic for 
Abrostola and Mouralia. 

171 (1-000) A dorsal patch of curved cornuti is synapomorphic for the members of clade 26. 

172 (0-500) The presence of two, short, straight cornuti on the vesica is apomorphic for clade 7, although 
there has been reversal in Chrysodeixis. In this genus, the position of the short cornuti is occupied by 
a series of large cornuti. Further studies are required to determine whether these two forms of cornuti 
are homologous. 

173 (1-000) A differentially sclerotized band of RHS sclerotization on the vesica is autapomorphic for 
the Argyrogrammatini. 

174 (1-000) A dorsal convex rounded plate on the vesica is synapomorphic for the members of clade 7. 

175 (0-333) Auricular cornuti have been convergently derived in Eosphoropteryx, Loboplusia and 
Macdunnoughia. 

Female abdomen A 1-7 

176 (0-333) A thickened pleuron on A7 is synapomorphic for the members of clade 31 (with an uncertain 
loss in Loboplusia - see Characters), but with a convergent derivation in Brachionycha. 

Ill (0-500) Concave lateral edges to T7 are also apomorphic for clade 3 1 (but with a subsequent reversal 
in Puriplusia), and thereby confirm the possible correlation with the previous character. 

178 (0-667) A pleural pouch is interpreted as being synapomorphic for the Plusiinae, Stictopterinae and 
Euteliinae, but with secondary loss in Magusa. The latter interpretation is supported by the presence 
in Magusa of a slight pouch, although this is not enlarged as in the three subfamilies. However, this 
conclusion must, for the present, be treated with caution for an alternative optimization of states 
interprets the pleural pouch as an independent derivation in the Plusiinae and in the Stictopterinae + 
Euteliinae. A small, concealed pleural pouch is autapomorphic for the Euchalciina. 

Anal papillae andA8-A9 + 10 intersegmental membrane 

179 (1-000) Flared anal papillae are synapomorphic for the Euteliinae and Stictopterinae. 

180 (0-500) The Stictopterinae + Euteliinae and Calophasia have convergently acquired densely setose 
inner surfaces to the anal papillae. 

181 (1-000) Frayed edges to the anal papillae are autapomorphic for the Stictopterinae. 

182 (1-000) A differentiated dorso-ventral row of setae on the anal papillae is an autapomorphy of the 
Plusiinae. 

183 (0-500) Short anal papillae characterize clade 22, but with reversal in Euchalcia. Several species of 
Euchalcia (e.g. E. italica (Staudinger), E. dorsiflava (Standfuss), E. augusta (Staudinger)) have even 



SYSTEMATICS OF THE PLUSIINAE 137 

more modified anal papillae, in which the apices are ventrally produced into acute points (Dufay, 
1968). Although the biology of these taxa is unknown (Dufay, 1968), similar modifications in the 
heliothine genus Schinia Hubner are associated with oviposition deep in flowers and unopened buds. 
Thus state in Euchalcia variabilis may merely represent an early stage in a reversal transformation 
series from short anal papillae to the highly-derived, blade-like forms. 

184 (0-500) The presence of an eversible dorsal pouch between the anal papillae and A8 can be 
interpreted either as autapomorphic for the Plusiinae, with reversal in the Abrostolini, or as 
independently derived in the Omorphinini and the Argyrogrammatini + Plusiini. 

185 (1-000) A sub-basal dorsal flange on the posterior apophysis is autapomorphic for clade 6. 

186 (1 -000) Thickened membrane between the LP V and the anal papillae is synapomorphic for Plusidia, 
Lamprotes and Pseitdeva. 

A8, ostium bursae & antrum 

187 (1-000) A subdorsal, longitudinal, desclerotized band on T8 is autapomorphic for the Abrostolini. 

188 ( 1 -000) A ventral pocket on the anterior edge of T8, dorsal to anterior apophysis, is also autapomor- 
phic for the Abrostolini. 

189 (0-250) T8 has become irregularly cut back antero-subventrally below the origin of the anterior 
apophysis independently in Anagrapha, clade 44 and clade 30 (but with reversal in Antoculeora). 

190 (0-500) Anterior apophyses have been independently lost in Lophoptera and the euteliines, or lost in 
the Stictopterinae + Euteliinae and regained in Stictoptera. 

191 (0-125) 'Protected' ostia bursae, as suspected (see Characters), have been acquired independently 
no less than eight times, in Brachionycha, Mouralia, Diachrysia, Rachiplusia, Antoculeora, Lam- 
protes, clade 32 and clade 44. When the true homologies of these structures have been determined, 
more information may be forthcoming but at present this character is useless. 

192 (1-000) Spinose pads lateral to the ostium bursae unite Macdunnoughia and Puriplusia, with the 
further development into recurved spinose grooves autapomorphic for the latter genus. 

193 (1000) Short, dense, upright scales near the mouth of the antrum are synapomorphic for the 
members of clade 30. 

194 (0-667) As initially suspected, setae along the inner ventral edge of the antrum (state 2) are 
convergently derived in Diloba and clade 26. Lamprotes is autapomorphic in that the setae are 
restricted to a subventral and lateral position (state 1). 

195 (0-500) A ventro-anterior extension of T8 shows the same distribution of states as character 190 and 
thus has the same two alternative sequences of state change. 

196 (0-167) An open, shallow antrum has been independently acquired on a total of six occasions, in 
Diloba, Cucullia, Chalcopasta, Agrapha, Pseudoplusia + Chrysodeixis and clade 21. A more 
thorough analysis of the homologies of the states of this character may reduce the homoplasy and 
allow more reliable information to be extracted from it. 

197 (0-500) An entirely membranous antrum has been convergently derived in Stictoptera and Cteno- 
plusia + Acanthoplusia. 

198 (1-000) State 1 of this character, in which the ventral edges of T8 are extended towards the midline, 
unites the members of clade 35. Further extension (state 2) is synapomorphic for the included genera 
Autoplusia and Antoculeora. State 3 (T8, antrum and LPV entirely fused ventrally) arises de novo 
from state in Rachiplusia. 

199 (1-000) The anterior apophysis arises level with the ventral edge of T8 only in Autoplusia and 
Antoculeora. 

200 (1-000) A heavily sclerotized, smooth, open antrum is also synapomorphic for Autoplusia and 
Antoculeora. 

201 (0-200) Thick, white membrane lateral to the ostium bursae has arisen five times independently in 
Plusiopalpa, Allagrapha, Lamprotes, Euchalcia and clade 44. 

202 (0-500) This character (T8 represented by a thin band dorsally) has the same distribution and choice 
of state change sequences as 190 and 195. 

Ductus bursae and corpus bursae 

203 (0-667) A long, narrow ductus bursae is interpreted as apomorphic for clade 9 but with a reversal in 
the included clade 14. Trichoplusia is autapomorphic for an extremely long ductus bursae (state 2). 
Given that the females of Plusiotricha are currently unknown, and both Trichoplusia and Thysano- 
plusia are heterogeneous for states 2 and 1 respectively, very little confidence can be placed in this 
character at present. 

204 (0-500) A corpus bursae that is produced at right angles to the long axis of the ductus bursae shows 



138 IAN J. KITCHING 

the same distribution and heterogeneity of states as the last, and is therefore subject to the same 
reservations. 

205 (0-200) Thickened white membrane on the corpus bursae at the entry point of the ductus bursae has 
been independently acquired by Cucullia, Trichoplusia, [Plusiotricha +] Eutheiaplusia and clade 24 
(but with subsequent loss in Lamprotes + Pseudeva). 

206 (0-500) An expanded portion of the ductus bursae with dorsally recurved edges has been con- 
vergently derived in Rachiplusia and Macdunnoughia + Puriplusia. 

207 (0-500) A short, lateral pouch on the ductus bursae occurs independently in Agrapha and Chry- 
sodeixis. 

208 (1-000) A small corpus bursae with highly elastic walls is synapomorphic for Diloba and 
Brachionycha. 

209 (0-182) The position of the origin of the ductus seminalis on the corpus bursae is highly homopla- 
sious and highlights the need alluded to above for a critical study of the homologies of the structures 
variously referred to as the fundus, apex and appendix bursae. The plesiomorphic condition appears 
to be an apical origin. Independent shifts to a fundular origin have occurred in Erythroplasia, 
Puriplusia, Loboplusia, Panchrysia, Oncocnemis, Cucullia and the Argyrogrammatini. Within the 
last, reversal has occurred in Pseudoplusia and clade 10, with re-reversal in [Plusiotricha +] 
Eutheiaplusia. However, at least two of these homoplasies (Puriplusia and Loboplusia) may be due 
to errors in deciding which end of the corpus bursae represents the fundus and which the apex (see 
also Ichinose, 1962b). For the present, character 209 is best dismissed as unreliable. 

210 (0-667) Regular ridges on the corpus bursae have been convergently derived in clade 47 and Cucullia 
+ Calophasia. Further modification to form a more or less circular pattern unites the Stictopterinae 
and Euteliinae. 

211 (0-667) A concave, circular signum unites the Stictopterinae and Euteliinae but is lost in Anuga and 
further modified into a ridge in Lophoptera. 

212 (1-000) A distinct differentiation of sclerotization of the corpus bursae granulations is synapomor- 
phic for Sclerogenia, Autoplusia and Antoculeora. 

213 (0-333) State 1 (granular ridges on the corpus bursae apex) is autapomorphic for Eosphoropteryx. 
Sclerotized ridges (state 2) either unite the members of the Autoplusiina, with subsequent loss in 
Erythroplasia, or are convergently derived in Rachiplusia and clade 31. Within the latter, very large, 
strong folds have either arisen in clade 31, only to be lost independently in Loboplusia and 
Allagrapha, or have been convergently acquired by clades 33 and 35. Only further examination of 
additional taxa will resolve these ambiguities but character 213 is undoubtedly important in assessing 
the interrelationships of the autoplusiine taxa. State 2 has also been convergently derived in Diloba, 
Chalcopasta, Omorphina and Plusiopalpa. 

214 (0-250) A recurved apex of the corpus bursae has been convergently derived in Pseudoplusia and 
clade 14. However, reversal is interpreted to have occurred in Acanthoplusia and Stigmoplusia. 

215 (0-429) This character was discussed in detail above in connection with the positions of Plusia, 
Cornutiplusia and Autographa. State 1 (a patch of thickened membrane on the RHS corpus bursae) is 
apomorphic for clade 40 but with a reversal in clade 44. State 2 (a shallow pouch) unites Autographa 
bractea and Cornutiplusia, while state 3 (a large pouch or appendix) is autapomorphic for the latter 
genus. State 3 arises de novo from state in Calophasia. 

Miscellaneous 

216 (0-500) Autoplusia and Rachiplusia are interpreted as being convergently sexually dimorphic. 

Clade analysis 

Each clade, as numbered in Figs 27, 28, is followed by its name (if applicable) and a list of its apomorphies 
in square brackets. The states printed in italics are those that are considered to be unique and unreversed 
synapomorphies. For those characters in which the state assignments to the internal nodes are ambiguous, 
the optimizations described in the preceding section are employed. It should be remembered, however, 
that these do represent only one of many possible interpretations. 

Clade 1: Plusiinae [15(1), 27(1), 29(7;, 37(7;, 32(0), 33(0), 35(0), 49(1), 131(1), 143(3), 782(7;, 184(1)]. 
The Plusiinae have been conventionally defined as the lashed-eyed, quadrifine noctuids. This study has 
demonstrated that this characterization is inadequate. Many of the outgroup taxa have lashed eyes 
(although it must be admitted that several were chosen for inclusion in this investigation expressly because 
they displayed this state), while one plusiine genus (Mouralia) has a distinctly trifine hindwing venation. 
Detailed study of the condition of hindwing vein M 2 showed that there are various degrees of reduction (see 



SYSTEMATICS OF THE PLUSIINAE 139 

character 15) and that a straightforward division into present and reduced/absent is too simplistic. Thus, a 
new characterization of the subfamily must be formulated. 

Of the 1 1 character states listed above, four are unique and unreversed for all members of the Plusiinae 
and support the monophyly of the subfamily. Thus, the Plusiinae can be defined, on present evidence, as 
those noctuids with a convex occiput (character 29); few apical styloconic sensilla on the proboscis (31), 
which has strengthening bars that are semicircular on the most apical portion (35); and anal papillae that 
bear differentiated, dorso-ventral rows of setae (182). 

With the possible exception of the last, however, these characters are not particularly useful as key 
characters. Several of the other seven homoplasious characters listed above are more suited to this role, 
providing that the exceptions are recognized and noted. Their inclusion results in a polythetic, but 
nevertheless workable, definition of the Plusiinae. The other characters are of three types. First, and most 
important, is character 184, the apomorphic state of which (the presence of an eversible dorsal pouch 
between the anal papillae and A8) is interpreted as characterizing the Plusiinae. Although it appears to 
have been lost in the Abrostolini, this pouch has not yet been found in any other group of noctuids and 
constitutes additional evidence corroborating the monophyly of the Plusiinae. Second, there are those in 
which the apomorphic state is found in all the plusiines but also occurs in one or more of the outgroup taxa: 
triangular pilifers (27; also found in Calophasia); a long, thin proboscis (32; also found in Calophasia + 
Cucullia), with smooth styloconic sensilla (33; also occurring in Cucullia); and a large thoracic crest (49; 
present also in Chalcopasta). Third, the apomorphic state may not only occur in the outgroups, but may 
also be transformed within the Plusiinae into other states, which themselves are found in the outgroups or 
are homoplasious within the Plusiinae. These are: a hindwing vein M 2 that is half as strong as M 3 (15; this 
state also occurs in Diloba, while within the plusiines, it is transformed into state 2 [a vestigial vein Af>] in 
Mouralia); the presence of saccular flanges (131; these are lost in the Abrostolini, Trichoplusia and 
Autoplusia); and a clavus borne on a peninsula on the sacculus ( 143; this condition is further modified in the 
Abrostolini and Anagrapha into state 2, in which the dorsal and ventral margins of the clavus are basally 
more or less level, convergent upon the condition found in Cucullia). 

Thus, it would appear that the key characters commonly used in the past are generally either too 
homoplasious (have too many exceptions to be of use to the non-specialist) or, on close examination, prove 
to have been misinterpreted. It is becoming increasingly apparent that satisfactory diagnoses of higher- 
level taxa will in future rely increasingly heavily upon character states that are not easily observable. This is 
a price that must be paid for a predictive and useful higher-level classification. 

The subfamily Plusiinae is represented throughout the World, with the exception of Antarctica and 
perhaps some of the more remote oceanic islands. A large number of species occur in Africa and the 
eastern Palaearctic, while, in comparison, the fauna of South America is relatively depauperate. Plusiines 
appear to be largely absent from tropical lowland forest habitats, while they reach their greatest diversity in 
tropical montane and temperate areas. Twelve species occur in Great Britain , either as residents or regular 
immigrants (e.g. Autographa gamma), while rare immigrants (e.g. Thysanoplusia orichalcea), singleton 
occurrences (e.g. Ctenoplusia accentifera Lefevbre) and doubtful natives (Euchalcia variabilis) account for 
a further twelve species. They rejoice under such common names as the Spectacle (Abrostola triplasia), the 
Beautiful Golden Y (Autographa pulchrina (Haworth)), the Burnished Brass (Diachrysia chrysitis) and 
the Gold Spot (Plusia festucae). Elsewhere in the World, the common names of plusiines are less 
imaginative, if perhaps more practical; for example, le Gamma (Autographa gamma), the Bean Leaf 
Skeletonizer (Autoplusia egena) and the Soybean Looper (Pseudoplusia includens). 

Few generalizations can be made regarding plusiine biology. The adults fly strongly and a number of 
species are migratory. However, they do not appear to be particularly attracted to light (Warnecke, 1953; 
Moucha & Novak, 1956; Eichlin & Cunningham, 1978). Thus a light-trap is neither an efficient nor 
accurate method for assessing plusiine population densities. Eichlin & Cunningham (1978) related an 
anecdotal observation that although numerous plusiines were observed feeding at night from flowers of a 
Button Bush (Cephalanthus occidentalis) that was growing less than 15 feet from a black light trap, only two 
moths actually came to the light. Many plusiines fly equally by day and by night. Silver Ys (Autographa 
gamma) are frequently seen in Britain on sunny late summer days feeding from Michaelmas Daisies (Aster 
spp.) in the company of butterflies and bees. 

As far as is known, temperate region species are univoltine, while in warmer climates, several 
generations per year seem likely. However, it should be stressed that only a minority of life histories, 
particularly of tropical species, has so far been elucidated. Multiple matings appear to be the rule, and the 
eggs are generally laid singly on the larval foodplant, probably between 200 and 300 per female (Eichlin & 
Cunningham, 1978). Plusiine larvae are usually green with white, often interrupted, longitudinal stripes. 
The head may bear a black, lateral stripe. Several species are sensitive to population density, becoming 
darker when overcrowded (Ichinose, 1959). There are generally five larval instars but a few species pass 



140 IAN J. KITCHING 

through six or even seven. In most species, the prolegs on segments A3 and A4 are reduced or missing (see 
below) and the larvae proceed by a looping action, thus giving rise to their common name of semiloopers. 
Pupation is usually in a cocoon spun between leaves. Pupal coloration has been shown to be temperature 
dependent in at least one species (Ichinose & Asawa, 1964) with darker pupae being formed at lower 
temperatures. According to Eichlin & Cunningham (1978), the prepupal stage lasts 1-2 days and the adults 
emerge about eight days after pupation. Little is known about plusiine courtship behaviour, but judging 
from the widespread occurrence of various tufts, brushes and pouches, pheromones produced by both 
sexes are likely to play an important role (see Gothilf & Shorey, 1976, for a comprehensive account of 
courtship in Trichoplusia ni) . 

Omorphinini [7(1), 20(0), 24(1), 42(1), 52(1), 59(1), 127(1), 137(1), 144(1), 147(1), 213(2)]. This tribe 
consists of a single genus and species, Omorphina aurantiaca. Of the character states listed above, ten are 
homoplasies occurring elsewhere in the Plusiinae or among the outgroups. This is only to be expected since 
the genus-level autapomorphies were largely eliminated from the data set prior to the analysis. However, 
characters 7 (yellow hindwings), 20 (randomly arranged scale bases), 24 (a prominent rim around the eye) 
and 137/147 in combination (oval valves with setae arising from marginal prominences) are useful 
recognition features. In addition, state 52(1), a truncate dorsal tegular arm, is autapomorphic. Although 
considered by Kostrowicki (1961) to be more correctly treated as an acontiine, the presence in Omorphina 
of the character states listed above for clade 1 argue strongly that Hampson (1913) was correct in placing 
the genus in the Plusiinae. 

Omorphina aurantiaca (Fig. 31) is a small moth, recorded only from Xizang Province, China (Tibet) and 
Sikkim, at around 15,000 ft (4500 m). Nothing is known about its biology except that the adult flies in July 
(Alpheraky, 1892). Omorphina has previously been likened to the Palaearctic amphipyrine Panemeria 
tenebrata (Scopoli) (Alpheraky , 1892) , but apart from the presence of marginal prominences on the valves, 
the genitalia of the two genera are dissimilar in both sexes. In particular, Panemeria lacks the row of setae 
on the anal papillae, while the cornuti on the vesica are very similar to those of Chalcopasta, which may 
indicate that the true affinities of Panemeria may lie with the Stiriini. 

Clade 2 [10(1), 28(0), 58(1), 73(1), 80(1), 99(1)]. This clade, which comprises all the Plusiinae except 
Omorphina (Omorphinini), is well supported by four unique and unreversed apomorphies: a dorsally 
delimited frons (28); an internal flange on the anterior edge of T2 that is produced subdorsally (73; see also 
clade 4); a particular form of St8 and ventral hair pencil (80), although this is frequently further modified; 
and the presence of a notch between the tegumen and uncus (99). In addition, all members of clade 2 
display a metallic sheen over much of the fore wings, although this is often weak and difficult to observe (10; 
the condition in Chalcopasta is almost certainly convergent but needs to be checked in other members of 
the Stiriini), and often bear a white spot on the fore tibia (58; this is frequently lost in the Plusiinae, occurs 
also in Chalcopasta and the Euteliinae, and is often impossible to score in many taxa due to additional 
pattern modifications). 

Clade 3: Abrostolini [1(1), 2(1), 51(1), 100(1), 102(1), 103(1), 107(1), 124(1), 125(1), 126(1), 131(0), 
134(1), 143(2), 169(1), 170(1), 184(0), 187(1), 188(1)]. There can be no doubt that the abrostoline genera 
Abrostola and Mouralia form amonophyleticgroup. Of the 18 character states listed above, no fewer than 
13 are unique and unreversed. Members of the tribe are best recognized by the form of the forewing 
stigmata (2), the fusion of the aedeagus to the juxta apex (124), the spatulate saccular lobe (134) and the 
desclerotized band and invaginated pocket laterally on T8 in the females (187/188 respectively). In 
addition, they are the only plusiines to possess raised scales in the forewing cell (1) and to lack an eversible 
pouch between the anal papillae and A8 in the females (184). Uniquely in the subfamily, as far as is known, 
the larvae of the Abrostolini bear prolegs, albeit somewhat reduced, on abdominal segments 3 and 4 
(Eichlin & Cunningham, 1978; see also below), although this condition is probably plesiomorphic. 

Abrostola [22(2), 121(2), 143(1), 169(2)]. This genus is probably paraphyletic relative to its sister-taxon 
Mouralia. The ventral hook on the vesica (169) and the heavily lashed eyes (22) appear to be restricted to 
A. triplasia (Fig. 32) and a few other, possibly closely related, species (e.g. A. kaszabi Dufay [Dufay, 1971]; 
A. trigemina (Werneberg)). The blunt, lobate VAVB (121) and the broad, rounded clavus (143) need to be 
confirmed in the other species of Abrostola. 

Abrostola includes about 35 species, of which four are North American and the remainder almost 
equally divided between Africa and the Palaearctic. None are known from South America, South East 
Asia or Australasia. Many species have only recently been described, particularly important contributions 
in this respect being those of Dufay (1957; 1958; 1960; 1971). The larvae of Abrostola are characteristic in 



SYSTEMATICS OF THE PLUSIINAE 141 

having a 'humped' segment A8, usually bearing two small, subdorsal points. In most species (e.g. A. 
triplasia, A. trigemina, A. major Dufay), there are diagonal, subdorsal white or cream stripes on segments 
Al-8, together with darker, more dorsal markings. On segments Al, A2 and A8, the latter fuse to form 
single, dorsal spots, which, together with the more pronounced pale markings, produce a very distinctive, 
somewhat eye-like, pattern. The overall colour can vary from green to violet-brown. The larva of A. 
asclepiadis is very different. A single, dried, inflated skin in the BMNH collection is uniform pale cream 
(?green in life), and is covered in a pattern of small, black spots, not all of which are associated with setal 
pinacula. The pattern is suggestive of an aposematic colour scheme, which is not unreasonable given the 
species' asclepiadaceous foodplant (see below). However, this hypothesis remains to be corroborated by 
the description or a colour photograph of a living larva. The biology of only a few species is known, 
primarily from the Holarctic fauna. Most species for which the larval hostplant is known feed on 
Urticaceae; usually Urtica itself (Crumb, 1956; Beck, 1960; Ichinose, 1962b; Bretherton et al., 1983; 
Miyata, 1983), but A. anophioides Moore utilizes Girardinia heterophylla (Gardner, 1947), while A. 
trigemina and A. major also feed on Boehmeria. A. abrostolina (Butler) is recorded from Fatoua 
(Moraceae) (Miyata, 1983), and other recorded foodplants include Epilobium, Lamium (Hampson, 1913) 
and Humulus (Bretherton et al., 1983). One species is exceptional, however, in that it has performed a 
remarkable hostplant shift; A. asclepiadis is the only plusiine known to feed exclusively on an asclepiad, 
Vincetoxicum hirundinaria (Hampson, 1913 as Cynanchum vincetoxicum; Beck, 1960; Weidlich, 1984 as 
Vincetoxicum officinale). Members of Abrostola are also unusual among the plusiines for overwintering in 
the pupal stage rather than as early instar larvae (Eichlin & Cunningham, 1978). The British species 
are known as 'spectacles' from the markings on the patagia, but this pattern is not found throughout the 
genus. 

Mouralia [14(2), 15(2), 18(2), 19(1), 64(3), 83(5), 136(1), 191(1)]. Mouralia currently includes only a single 
described species, M. tinctoides (Fig. 33). Of the above eight listed character states, three constitute good 
autapomorphies for the genus: the presence of spines on the inner surface of the hind tibia (64); the 
extremely long sternal hair pencil , arising from a single pocket that almost reaches the base of the abdomen 
when retracted (83); and the presence of basally-directed scales on the valve apex (136). Although this 
latter condition was coded as also present in the Argyrogrammatini (q.v. ), the form of the scales differs and 
so the two conditions probably represent distinct characters. In the Argyrogrammatini, the scales are 
simple and setose, often quite stiff and usually easily detached, while in Mouralia. these scales are long, 
soft, curved, apically spatulate and persistent (Fig. 348). In addition, Mouralia is the only plusiine with a 
trifine hindwing venation (15) and has a female frenulum reduced to two setae (14; a condition also found in 
clade 6, Rachiplusia, Autoplusia and Brachionycha). 

There are also numerous abdominal and genitalic autapomorphies that were not included in the analysis. 
These include: a long, thin, medial apodeme projecting anteriorly from the anterior margin of St7 in the 
male (this may be functionally correlated with the long hair pencil pocket); an apically, laterally flattened 
uncus (Fig. 278); a pair of spatulate flaps arising from the dorsal sclerotized band on the anal tube; an 
ampulla (Fig. 348); valves thickly covered ventrally and exteriorly with long, flexuous, pale yellow, 'sticky' 
(in the sense that they adhere to forceps) scales; valves in which the apices turn through 90 degrees so that 
the costal edges become internal and the saccular edges become external (Fig. 348); and a corpus bursae 
with a very elongate, tapering fundus that broadens to a blunt tip. 

A second undescribed species of Mouralia has recently been discovered in Bolivia (F. Navarro, pers. 
comm.). It differs in several features from M. tinctoides. The most notable of these are (as assessed from a 
photographic slide of a male genitalic preparation) the presence of only a rudimentary hair pencil and a 
valve apex that lacks all the modifications of M. tinctoides, simply tapering to a point as in many abrostolas. 
Thus this new species may represent an intermediate form between the genus Abrostola and Mouralia 
tinctoides. 

Another candidate for such an intermediate taxon is Abrostola anophioides. This is a large species found 
in the Himalayan foothills from Uttar Pradesh in NW. India to Sichuan Province, China. Unlike other 
abrostolas, the males of A anophioides have a short, single hair pencil, while the fundus of the corpus 
bursae is elongate and apically swollen as in Mouralia tinctoides. A. anophioides and the new Mouralia 
species strongly suggest that one result of a detailed study of the Abrostolini might be the synonymy of 
Mouralia with Abrostola. 

Mouralia tinctoides is widespread throughout the Neotropics, penetrating as far north as the SW. United 
States. Its life history has been described in detail by Comstock (1937). The general appearance of the larva 
is that of an Abrostola but it differs markedly in hostplant , feeding on Tradescantiafiuminensis and Zebrina 
pendula (Commelinaceae) (Eichlin & Cunningham, 1978) (although the latter may be an error; Eichlin, 
1975). Unusually for the Plusiinae, there are seven larval instars (the norm is five). 



142 IAN J. KITCHING 

Clade 4 [3(1), 4(1), 6(1), 59(1), 73(2), 83(3), 106(1), 157(2), 158(1), 162(1)]. The Argyrogrammatini and 
the Plusiini are united as sister-groups primarily on the basis of two synapomorphies: a distinct, bulbous 
coecum (157) and a unique form of the internal flange on the anterior edge of T2, in which the two 
subdorsal lobes (see clade 2) are fused medially but with a medial ventral emargination (73). In addition, 
several character states occur only in clade 4, although in not all of the constituent genera; these are the 
characteristic plusiine Y-mark (3), a pink terminal line on the forewing (4, see also clade 25), a scale tooth 
(6), restricted sclerotization on the aedeagus (158) and a subapical cornutus (162). Further, in all members 
of the Argyrogrammatini and Plusiini, the LASTs are fused around the base of the uncus, although this 
condition recurs in Chalcopasta. 

As far as is known, all members of clade 4 that hibernate do so as early instar larvae. Also, the prolegs are 
vestigial or absent on segments A3 and A4 in all larval instars (but see Anadevidia). 

Clade 5: Argyrogrammatini [14(1), 18(0), 89(1), 108(1), 136(1), 144(1), 158(2), 173(1), 209(1)]. This 
primarily tropical tribe of plusiines is quite well characterized by three synapomorphies. All argyrogram- 
matines have secondary hair brushes arising on St8 (89), basally-directed setose scales apically on the valve 
(136, see also Mouralia), and a differentially sclerotized band on the vesica (173). Seta 1 of the female 
frenulum is also reduced (14; see Mouralia, Autoplusia, Rachiplusia) and the ventral sclerotization of the 
aedeagus is further restricted to a distinct narrow band (158; with convergence occurring in the autoplu- 
siine genera Erythroplasia and Macdunnoughia) . 

Anadevidia [3(0), 6(1), 105(1), 116(1), 148(0)]. Anadevidia is an enigmatic genus, of somewhat uncertain 
position. Both Kostrowicki (1961) and Ichinose (1962b) refrained from making any explicit statements as 
to its relationships, although the latter derived the larval setal pattern from that of Thysanoplusia (as 
Diachrysia), a relationship formalized by Chou & Lu (1979) as the 'Plusiini'. Kljutshko (1985b) placed 
Anadevidia in the Plusiini, where it remained after the reanalysis performed above, on the basis of lack of 
prolegs in the mature larva and setae SD1 on segment A9 being 'normal'. This similarity to members of the 
Plusiini is supported further by character 116, in that both taxa possess a strongly concave pleurite. 
However, as described above, there is stronger evidence (characters 89, 136 and 173) to suggest that 
Anadevidia correctly belongs in the Argyrogrammatini. 

It may yet be shown that Anadevidia is in some way related to Chrysodeixis (+ Pseudoplusia). Both 
genera bear spatulate scales on segment A8 in the male (although these are interpreted here as 
independent conditions). Also, the vincular arms in Anadevidia are long and narrow, and superficially 
resemble those of Chrysodeixis and Pseudoplusia, although they are separate in the former genus and 
intimately fused into a thin spine in the latter two. 

Anadevidia currently contains only two species, A. peponis (Fig. 34) and A. hebetata. The former is 
widespread throughout the Indo-Australian region, while the latter is apparently restricted to India and 
Japan. In general appearance, they are dull grey-brown moths with no distinguishing pattern features, and 
bear a superficial resemblance to species of the genera Allagrapha and Diachrysia (especially D. leonina 
and D. bieti (Oberthiir)). The larvae of Anadevidia have several unusual features. They are the only 
plusiines in which the first instar larvae possess on segments A3 and A4 well developed prolegs that are 
subsequently lost completely in the later instars. The larvae also bear enlarged pinacula on some segments, 
giving them a warty appearance. Anadevidia is oligophagous, feeding on members of the Cucurbitaceae, 
although A. peponis has been recorded from Mercurialis (Euphorbiaceae) and Paulownia (Scrophular- 
iaceae) (Miyata, 1983). The larval chaetotaxy of Anadevidia has been described by Ichinose (1958b; 
1962a; b). 

Clade 6 [12(1), 14(2), 76(1), 77(1), 83(2), 91(1), 94(2), 110(1), 167(1), 185(1)]. Three very good 
synapomorphies characterize clade 6: the form of St7 in the male (76) ; the presence of an invaginated sac in 
the vesica (167); and the presence of a sub-basal dorsal flange on the posterior apophysis (185). The males 
of the majority of the constituent taxa also bear dorso-lateral hair tufts on at least segments A5 and A6 (77, 
except Zonoplusia and Pseudoplusia), have dorsally twisted lateral arms of St8 (91, except Pseudoplusia) 
and have broad, 'greasy' scales apically on the dorsal arms of St8 (94, again, except Pseudoplusia). Overall, 
clade 6 is very well supported, allowing for the multiple reversals of Pseudoplusia (q.v.). 

Clade7[93(l), 105(1), 108(0), 128(1), 162(0), 172(1), 174(1)]. The three genera ofthis clade are united bya 
single synapomorphy, a dorsal, convex, rounded plate on the vesica basally (174). In addition, a pair of 
short, straight cornuti occur in both Dactyloplusia and Pseudoplusia, but not in Chrysodeixis (172). 
However, as discussed above (Character analysis), the condition in the latter genus may prove to be 
derivable from that in the former two. Also, all three genera have a juxta that is apically scobinate (128), 
although this condition recurs in Thysanoplusia, Plusiopalpa and Cucullia. 



SYSTEMATICS OF THE PLUSIINAE 143 

Dactyloplusia [60(1), 78(1), 80(2), 92(1), 110(0), 149(1)]. In the present analysis, Dactyloplusia is 
interpreted as being the sister-group of Chrysodeixis + Pseudoplusia, on the basis of the characters listed 
above forclade7. However, there are an almost equal number of character states that suggest Dactyloplu- 
sia may instead be related to clade 10, and in particular, Thysanoplusia. These include the presence of a 
hind basitarsal comb (60) , latero-ventral flaps on the anterior bar of St8 (80) , a sub-basal lobe on the lateral 
arms of St8 (92), a sharply hooked harpe (149), as well as the apically scobinate juxta referred to in the 
preceding paragraph. Dactyloplusia is also autapomorphic in having the dorso-lateral hair tufts on segment 
7 enclosed in a groove (78). 

Dactyloplusia currently contains only a single species, D. impulsa (Fig. 50). Holloway (1985) gives the 
distribution as Sri Lanka, India, S. China, Sundaland, Sulawesi and New Guinea. However, by far the 
majority of specimens in the BMNH collection were caught in Sri Lanka. Nothing is known of the biology 
of D. impulsa. Holloway (1985) captured a Bornean specimen 'on a river bank with disturbed vegetation 
and alluvial forest near a limestone cliff at 100 m' . 

Clade 8 [11(1), 111(1), 120(1), 130(0), 144(0), 196(1)]. A sister-group relationship between Pseudoplusia 
and Chrysodeixis is hypothesised on the basis of two synapomorphies: a long, thin, spine-like vinculum 
(111; the condition found in Stigmoplusia is considered to be non-homologous) and a very narrow, almost 
straight pleurite (120). Ichinose (1973) also recognized the great similarity between these two genera and 
thought that Pseudoplusia might eventually prove to be best placed in Chrysodeixis, perhaps as a subgenus. 

Pseudoplusia [59(0), 77(0), 83(0), 91(0), 93(0), 94(0), 725(2), 209(0), 214(1)]. Pseudoplusia is perhaps 
most notable for the total reduction of the male scent organs, both those of St8 and the dorso-lateral hair 
tufts (characters 77, 83, 91, 93, 94). This may prove to be due to the geographic isolation (one species in the 
Americas, the other on Saint Helena) from other members of clade 7, resulting in a relaxation of the 
selective pressures that maintained the elaborate scent organs, which were thus lost. Pseudoplusia also 
displays a number of autapomorphies. The males possess a hook-like process near the base of the valve 
costa, an extremely long, thin, sinuous harpe, and bear a sharp spine medially on the posterior margin of 
the juxta (Fig. 341; similar to the type of spine found in some species of Syngrapha). The juxta is also 
latero-apically spinose (128). In the females, the apex of the corpus bursae is extremely elongate and 
tapered, projecting first anteriorly and then recurving posteriorly. 

The genus Pseudoplusia contains two species, P. includens (Fig. 38) and P. dalei (Wollaston) comb. n. 
(see Appendix 4). The former is widespread from Canada to Argentina and Chile (Forbes, 1954; Angulo & 
Weigart, 1975) and is an important pest of many cultivated plants (Canerday & Arant, 1967). It is 
commonly known as the Soybean Looper (Eichlin & Cunningham, 1978) or, less frequently, the False 
Cabbage Looper (Canerday & Arant, 1967). An icosahedral virus has recently been isolated from P. 
includens (Chao, Young & Kim, 1984) and may prove useful as a biological control agent. P. dalei is 
endemic to the mid-Atlantic island of Saint Helena. Little is known of its biology except that the adults 
have been seen frequenting the flowers of geraniums and Pittosporum and that it may belong to the 
nearly-extinct scrubland fauna of the island (Wollaston, 1879). 

Chrysodeixis [12(0), 77(3), 83(5), 94(1), 97(1), 105(0), 127(1), 172(0), 207(1)].. In contrast to its sister- 
genus, Chrysodeixis is most notable for the often extreme elaboration of the scent organs on the male 
abdomen. The progressive reduction of the sternal hair pencils, with the concomitant modification of the 
secondary hair brushes, T8 and St8, has already been described above (Terminology). In addition, the 
more derived species, such as C. chalcites (Fig. 39), have distinct additional dorso-lateral hair tufts on 
segment A4 (77); broad, spatulate scales apically on the dorsal arms of St8 (94); and an additional two pairs 
of hair pencils arising subventrally from discrete pockets in the membrane between segments 7 and 8. In the 
more plesiomorphic species, C. argentifera (Fig. 40), the sternal hair pencils are more fully developed but 
arise from a single pocket (83), much as in Mouralia. 

Chrysodeixis is a moderate-sized genus of about 25 species found throughout the Old World tropics. 
However, by far the majority of species occur in SE. Asia and Australasia. Most are relatively restricted in 
range but three, presumably by virtue of a strong migratory habit, have become very widespread. C. acuta 
is found more or less throughout the entire range of the genus (Dufay, 1970a; Pinhey, 1975). It has even 
been recorded from Great Britain despite apparently never having been captured on mainland Europe 
(Bretherton etal. , 1983). C. chalcites is sympatric with C. acuta over the western part of the latter's range, 
but is replaced by C. eriosoma from approximately Indonesia eastwards. There is some debate as to 
whether C. chalcites and C. eriosoma are (Pinhey, 1975) or are not (Kostrowicki, 1961; Dufay, 1970a) 
conspecific. The biology of the two species is very similar and although allopatric C. eriosoma can be 
distinguished from C. chalcites by differences in the cornuti (Kostrowicki, 1961), in the area where the two 



144 IAN J. KITCHING 

species are thought to overlap, the distinction is not as clear cut and there is confusion as to their identities 
(Holloway, 1985). C. chalcites has also been taken in Great Britain and, in contrast to C. acuta, is 
frequently recorded from Mediterranean Europe (Bretherton etal. , 1983). 

One section of Chrysodeixis, the minutus Dufay species-group (Holloway, 1985), does not show the 
needle-like vinculum but does possess additional hair pencils between segments A7 and A8. The 
relationships of these species need to be assessed carefully in any future analysis of the interrelationships of 
the members of clade 8. 

Clade 9 [59(0), 203(1), 204(1)]. This clade is not supported by any good synapomorphies and will probably 
not be corroborated by future studies. Two of the character states, an elongate ductus bursae (203) and a 
transversely orientated corpus bursae (204), only occur in the included genera Zonoplusia, Thysanoplusia, 
Trichoplusia and Eutheiaplusia (with Plusiotricha unknown), with both states being subsequently lost in 
clade 14. However, the apomorphic states for 203 and 204 are not found throughout Trichoplusia and 
Thysanoplusia (Dufay, 1970a) {Zonoplusia and Eutheiaplusia are treated here as monotypic) and it is thus 
highly unlikely that they will be shown in future to support a monophyletic group of genera comprising 
clade 9 less clade 14. Certainly, before such a study can be attempted, a critical reappraisal of Trichoplusia 
and Thysanoplusia must be undertaken (see also Plusiotricha). 

Zonoplusia [4(0), 58(0), 77(0)]. Zonoplusia is treated here as a monotypic genus, comprising only the type 
species, Z. ochreata. When originally erected (Chou & Lu, 1978a), five species were placed in Zonoplusia. 
Two were unavailable for examination (Z. longisigna (Chou & Lu) and Z. brevistriata (Chou & Lu)) but 
are probably closely related to, if not synonymous with, either of the other two species. These are 
albostriata, a current member of Ctenoplusia (for the moment, see 60 and 133) and daubei, presently placed 
in Thysanoplusia. Ichinose (1973) included Z. ochreata in Thysanoplusia, although it differed from T. 
intermixta and T. daubei in the larvae having very fine cuticular ornamentation (like Chrysodeixis, 
Trichoplusia and Ctenoplusia s.l.) rather than large and hair-like microtrichia. 

Z. ochreata (Fig. 49) is a small moth, in which the Y-mark has been modified into an oblique, pale grey, 
double line. The subapical cornutus is unusual in that it arises from a sclerotized base that is almost as long 
as the cornutus, thus giving the appearance of a double cornutus. Z. ochreata is found in the eastern 
Palaearctic and throughout the Oriental tropics and has also been recorded from Queensland, Australia 
(although otherwise not east of Sundaland and the Philippines (Kostrowicki, 1961; Holloway, 1985)). The 
only known larval foodplant is Polygonum thunbergii (Polygonaceae) (Miyata, 1983). 

Clade 10 [80(2), 84(1), 92(1), 93(2), 209(0)]. This clade is also not supported by any good synapomorphies. 
As was discussed above, states 80(2) and 92(1) are shared with Dactyloplusia, which may have its true 
affinities with members of clade 10. Sclerotized bars on the sternal hair pencil pockets (84) are restricted to 
members of this clade but are absent in Thysanoplusia, Plusiopalpa and Acanthoplusia. Similarly, dorsal 
arms of St8 that are apically fused into a rounded point (93 : 2) are found only in clade 10 , although this state 
is further modified in clade 18 (93:3) and Thysanoplusia (93:1, convergent upon the condition found in 
Chrysodeixis and Dactyloplusia). 

Clade 11: Trichoplusia s.l. [60(1), 83(3), 144(0)]. No unique and unreversed synapomorphies have been 
found to link Trichoplusia and Thysanoplusia as sister-groups. The form of the hind basitarsal comb (60), 
sternal hair pencils (83) and clavus (144) are all probably the plesiomorphic conditions for these characters 
in the Plusiinae (whether by primary symplesiomorphy or by secondary reversal). 

Trichoplusia s.str. [105(1), 110(0), 131(0), 149(1), 203(2), 205(1)]. Trichoplusia s.str. (i.e. sensuMcDun- 
nough, 1944; Ichinose, 1973) is an extremely heterogeneous genus and appears to have been adopted as the 
'dustbin' of the Argyrogrammatini, in much the same way as 'Plusia' s.l. accommodates any generally 
unplaceable plusiine. As a result, the genus currently contains about 50 species. The remarks made below 
thus properly refer only to T. ni. 

The only good autapomorphy for T. ni (Fig. 43) discovered in this study was the extremely long ductus 
bursae (203), which was the main character state emphasised by McDunnough (1944) when he originally 
proposed the genus. A similar long ductus bursae also occurs, however, in T. oxy gramma (McDunnough, 
1944; Eichlin & Cunningham, 1978) and in T. gorilla (Holland) (Dufay, 1970a), where it is even longer, 
being some 3-4 times the length of the body. T. ni is the only argyrogrammatine examined to lack a 
saccular flange (131) and, like Dactyloplusia and Agrapha, possesses a sharp, hooked harpe (149). 

Trichoplusia is badly in need of a critical and thorough revision. Species have been added somewhat 
casually since Dufay (1970a) began the rapid expansion of the genus. It is almost certainly polyphyletic with 



SYSTEMATICS OF THE PLUSIINAE 145 

regard to at least Thysanoplusia, Dactyloplusia, Zonoplusia and Ctenoplusia. This much is immediately 
obvious from the length of the couplet (24 lines) necessary to distinguish Trichoplusia from Ctenoplusia in 
Dufay's (1970a) key. A flavour for the problem can be gained from the description in the key of the form of 
the cornuti in Trichoplusia (translated freely) : 'aedeagus often armed with a small , spiniform or needle-like 
cornutus, accompanied or not by a mass of fine spicules, or furnished with a large number of spines, or only 
with a more or less dense edging of denticles; rarely almost unadorned'. The corresponding 'condition' in 
Ctenoplusia reads: 'aedeagus armed with many cornuti or with a large and long, stick-like cornutus, never 
with a single, fine, spiniform cornutus, sometimes with several small cornuti'. Trichoplusia almost certainly 
contains a number of monophyletic units and their elucidation is probably the most pressing, yet most 
difficult, task in plusiine systematics. 

T. ni is the world's most widespread plusiine. As race brassicae (Riley), it occurs from Canada to 
Central, and even South, America (Kostrowicki, 1961; Eichlin & Cunningham, 1978) and had been 
introduced into Hawaii by 1939 (Zimmerman, 1958). As race ni, the species occurs throughout the 
southern Palaearctic, Africa and most of the Oriental and Australasian regions, reaching South Africa, 
Madagascar, Korea and New Zealand. The two races are very dubiously distinct (the forewings of ni are 
said to be lighter and often greyer than those of brassicae) and the separation is often not made (e.g. Eichlin 
& Cunningham, 1978; Franclemont & Todd, 1983). 

Commonly referred to as the Cabbage Looper or the Ni Moth , T. ni is a very important pest of numerous 
crop plants, but particularly of brassicas (as one of the common names implies). The list of its larval 
foodplants is impressive. Eichlin & Cunningham (1978) list over 60 genera from a wide range of families 
and the species would appear to be able to eat virtually anything, including plants with apparently 
considerable chemical defences (e.g. Asclepias, Crotalaria, Euphorbia and Senecio). I reared large 
numbers, from New Zealand stock, in the BMNH and the only plant that was actively refused was the 
introduced Japanese knotweed, Reynoutria japonica. Because of its pest status, T. ni has been the subject 
of much research activity aimed at its control or destruction and this in turn has generated an enormous 
amount of literature on the subject (see any appropriate abstracting journal). 

Thysanoplusia [2(0), 9(1), 84(0), 93(1), 112(1), 128(1), 159(1)]. Thysanoplusia was originally erected by 
Ichinose (1973) to accommodate those species previously incorrectly placed in Diachrysia, due to 
confusion over type species (see Review). The genus originally included T. intermixta (Fig. 47), T. 
orichalcea, T. daubei (Fig. 48) and T. (now Zonoplusia) ochreata. Suggested future candidates for 
inclusion were florina, homoia (Dufay), ignicollis (Dufay), semirosea (Walker) and viettei. Although 
stated to be separable on the basis of larval and adult characters, the latter were never clearly stated. Thus, 
the characterization of Thysanoplusia came to depend upon larval features. Most notably, Thysanoplusia 
differed in having a very slender seta SD1 on segment A9 and in bearing setae SV1 and SV2 on A2 on a 
single pinaculum (Ichinose, 19626; 1973). The genus was retained, with reservations, by Holloway (1985), 
who added T. ekekei (Bethune-Baker) and T. bipartita (Snellen). However, most authors treated 
Thysanoplusia as synonymous with Trichoplusia (e.g. Dufay, 1975; 19826) and eventually it was so treated 
in the Japanese checklist (Inoue etal., 1982). Although I accept that Thysanoplusia defined in terms of the 
two larval characters may prove to be a monophyletic unit, until such time as good adult synapomorphies 
are found (or, alternatively, and far less likely, the larvae of the outstanding species are described), the 
concept is of limited value. Thysanoplusia is best either sunk completely into Trichoplusia until a 
comprehensive revision can be undertaken, or restricted to a group of species centred upon T. intermixta 
(perhaps including T. daubei and its relatives), but always remembering that this is at best an incomplete 
group. 

T. intermixta is a member of that subsection of Thysanoplusia with a conspicuous metallic golden-green 
patch on the forewing (9). Five species are currently recognized in this species-group: T. ekekei from New 
Guinea; T. bipartita from Sulawesi and Borneo (both these species have diffuse patches); T. intermixta 
from the Oriental tropics as far as Java and Japan (Holloway, 1985); T. florina, a Madagascan endemic; 
and T. orichalcea, which is widespread in the Old World tropics and subtropics. Like other highly 
successful and widespread plusiines (Trichoplusia ni, Chrysodeixis acuta, C. chalcites, Pseudoplusia 
includens, Autographa gamma), Thysanoplusia orichalcea is strongly migratory. It is occasionally taken in 
Great Britain and Scandinavia and has recently become established in New Zealand (1984), possibly from 
New Caledonia or Australia. Again, like the other successful species, T. orichalcea is a serious agricultural 
pest throughout much of its range, including New Zealand (J. Dugdale, pers. comm.), where it attained 
this status within a mere 12 months of its recorded arrival. 

Clade 12 [12(0), 77(2), 91(2), 138(1), 162(0)]. This clade is another in the Argyrogrammatini that is not 
corroborated by any good synapomorphies. Two features are restricted to members of this clade: the 



146 IAN J. KITCHING 

presence of weak dorso-lateral hair tufts on segments (A2-)A4 in the male (77) and lateral arms of St8 that 
are twisted dorsally with a deep, basal concavity (91). However, both of these states are later lost, in clade 
16 and Plusiopalpa respectively. 

Clade 13 [108(0), 140(1), 159(1), 160(1), 205(1), 209(1)]. Eutheiaplusia and Plusiotricha are considered to 
form a sister-group pair on the basis of two character states: abroad, blunt point on the sacculus (140) and 
the characteristic serpentine form of the inverted vesica (160) . In addition , the ornamentation of the vesica 
is unusual in comprising more or less uniformly distributed small spines ( 159) , a condition that is elsewhere 
only found in Thysanoplusia. 

Plusiotricha [11(1), 60(2), 61(1), 112(3), 119(1), 133(1), 144(2)]. As discussed above, Plusiotricha shares 
with Ctenoplusia and Acanthoplusia the unique blade-like setae along the sacculus. In addition, Plusio- 
tricha also shares another state with clade 16, a pleurite and VAVB fused for about half the length of the 
former (119), and has a similar form of basitarsal comb (60) to Ctenoplusia. These character states argue 
strongly that a future, more thorough study may show these three genera (and Eutheiaplusia, see below) 
form a monophyletic unit. Plusiotricha is unique in having hook-like projections on the vinculum arms 
towards their apices (112). 

Of more immediately importance would be the discovery and examination of a female of Plusiotricha. 
The genus currently contains three species, P. livida, P. carcassoni Dufay and P. fletcheri Dufay, 
represented in collections by seven, one and ten males respectively. A further four males are known that 
may represent a fourth species (Fig. 51). Female specimens are urgently needed to check whether the 
predicted female character states, particularly the elongate ductus bursae (203), found in Eutheiaplusia do 
also occur in Plusiotricha. 

Nothing is known about the biology of Plusiotricha. The holotype of Plusia dyscapna Fletcher (a 
synonym of Plusiotricha livida) includes a pupal exuvium and final instar larval skin, but no life history 
information accompanies the specimen. Neither was any mention made of the immature stage material in 
the original description (Fletcher, 1963). The genus Plusiotricha appears to live in mountainous areas at an 
altitude of about 4000-5000 ft (1200-1500 m) in the tropical rainforest belt of Africa, from the Ivory Coast 
to Kenya and Zimbabwe (Dufay, 1972). The largest number of specimens has been taken in Tanzania. 

Eutheiaplusia [4(0), 83(0)]. Eutheiaplusia can be characterized as a Plusiotricha in which the male scent 
organs have become highly reduced and presumably non-functional (83; and thus sits in relation to 
Plusiotricha as Pseudoplusia does to Chrysodeixis), and in which the blade-like saccular setae are replaced 
by fine, somewhat flattened, upcurved hairs and the vincular hooks are lacking. It could thus represent 
either the sister-group of Plusiotricha or may eventually prove to be the most basal lineage of a clade 
comprising the present clades 13 and 16. The vesica of the male illustrated by Dufay (1970a) bears a long, 
thin, subapical cornutus absent in the specimen examined in this study. The presence or absence of this 
structure in E. pratti (Fig. 52) requires confirmation. 

Eutheiaplusia comprises a single species, E. pratti, endemic to Madagascar. Nothing is known of its 
biology, although the species appears to be quite widespread on the island in forest at altitudes of between 
730 and 1700 m (Dufay, 1970a). 

Clade 14 [86(1), 87(1), 112(1), 203(0), 204(0), 214(1)]. As was found for clades 9, 10 and 12, there are no 
good synapomorphies for clade 14. Accessory pouches in the sternal hair pencil pockets (86) occur only in 
this clade (but not in Ctenoplusia or Plusiopalpa), as do sternal hair pencils in which the outer scales are 
shorter than the inner (87; but not in the included clade 17). Dorsal emarginations of the vincular arms 
(112) do characterize the members of this clade but have been convergently acquired by Thysanoplusia. 

Agrapha[6(l), 12(1), 79(1), 88(1), 98(1), 105(1), 130(0), 144(2), 149(1), 162(1), 196(1), 207(1)]. Agrapha 
currently contains three described species: A. ahenea, A. meretricia (Schaus) comb. n. and A. calceolaris 
(Walker) comb. n. There are also two undescribed species that were allocated manuscript names by E. L. 
Todd (note in BMNH collection, 1972). One, here termed species A, is represented in the BMNH 
collection by seven specimens from Colombia, Bolivia and Peru, and includes noctuid genitalia prepara- 
tion 5193 (male). The second species (sp. B) is unrepresented in the BMNH collection, is of unknown 
provenance and depository, and was not examined. 

Of the included species, three (A ahenea (Fig. 57), A. meretricia and sp. A) present a distinctive facies, 
particularly with regard to the highly developed dorso-lateral hair tufts, which give the abdomen a 
truncate, bushy appearance. The same three species (A. calceolaris was not examined) have unique 
flattened scales on the valve (see 133, Character analysis) and A. ahenea (and maybe A. meretricia and sp. 



SYSTEMATICS OF THE PLUSIINAE 147 

A) is the only argyrogrammatine to have discrete edges to the dorso-lateral hair tufts on both segments A5 
andA6(79). 

Agrapha [ahenea] shares two character states with Argyrogramma (a distally dilated uncus, 98, and a 
long inner tuft in the sternal hair pencil, 88) that must be interpreted as convergent. The hook-like harpe 
(149), referred to above in connection with Dactyloplusia and Trichoplusia, is present in A. ahenea and A. 
meretricia but in sp. A, the harpe is 'normal' (state 0) but with a small apical point. All species of Agrapha, 
with the possible exception oiA. calceolaris, have very long antennae (although this is difficult to quantify). 
I consider the recent inclusion of Trichoplusia oxygramma in Agrapha by Franclemont & Todd ( 1983) to be 
incorrect as this species has none of the autapomorphies of Agrapha just enumerated. Similarly, I reject 
their synonymy of Ctenoplusia and Acanthoplusia with Agrapha. No evidence was found in this study to 
suggest that these three genera form a monophyletic unit. 

Agrapha is the only argyrogrammatine genus wholly endemic to the New World, being found from Costa 
Rica to Peru. A. calceolaris and a possibly undescribed species (not sp. A, may or may not be sp. B) occur 
on Jamaica. Nothing is presently known about the biology of any of the species but they appear to be 
widespread through a range of altitudes. A. meretricia is recorded from 320 ft (100 m) in Colombia to 
6500+ ft (2000+ m) in Peru. Most specimens in the BMNH collection have been taken above 2500 ft 
(750 m). 

Clade 15 [92(3), 144(0)]. The five genera of this clade are united solely on the basis of their possessing an 
attenuate and dorsally recurved sub-basal lobe on the lateral arms of St8 (92). The degree of development 
of the clavus (144) has already been shown to be a worthless character (see Character analysis). 

Clade 16: Ctenoplusia s.l. [4(2), 60(1), 77(1), 119(1), 133(1), 197(1)]. Given that blade-like saccular setae 
(133) and a pleurite partially fused with the VAVB (119) are interpreted as convergently derived in this 
clade and in Plusiotricha, only two good synapomorphies were found to unite Ctenoplusia and Acanthoplu- 
sia: the tendency for the pink forewing terminal line to be concentrated at the apex of vein My (4) and the 
totally desclerotized, membranous antrum (197). The former is by no means universal in Ctenoplusia but 
does occur (or its more derived state 3) in C. limbirena, C. amydra Dufay, C. accentifera and the C. 
furcifera (Walker) species-complex. This corroborates the hypothesis that Acanthoplusia is the sister- 
group of a section of Ctenoplusia and supports Ichinose's (1973) synonymy. The occurrence of the 
desclerotized antrum in other species of Ctenoplusia and Acanthoplusia needs to be confirmed. 

Acanthoplusia [83(3), 84(0), 108(0), 214(0)]. Acanthoplusia was erected by Dufay (19706) to accommo- 
date five species: A. tarassota (Fig. 53) (NE. Himalaya, Java, Bali), A. ichinosei (Dufay) (Japan), A. 
agnata (China, Japan), A. latistigma (Prout) (Ceram) and A. eugrapha (Hampson) (New Guinea). 
Subsequently, other species were included (Dufay, 1910b; 1974; 1982): A. adiaphora Dufay (Taiwan), A. 
herbuloti Dufay (Luzon, Philippines), A. sigillata Dufay (Luzon, Philippines) and A. vermiculata (Sumat- 
ra). These species differed from Ctenoplusia in the number of cornuti, the shape of the male tergites of 
segments A5 and A6, and the presence of a pad of inflated scales on the outside of the valve. The first two 
characters were found to be too variable within the Plusiinae as a whole to be satisfactorily coded in the 
present study but may have value in a species-level investigation. The scale pad is, however, a good and 
easily recognizable apomorphy for Acanthoplusia, which is doubtless a monophyletic taxon, even if not 
finally deserving full generic status. 

Three species are known from immature stages: A. tarassota, A. ichinosei and A. agnata. All three 
appear to be somewhat polyphagous (Ichinose, 19626; Miyata, 1983). The larvae of A. agnata are unusual 
in the Argyrogrammatini in completely lacking prolegs on segments A3 and A4, instead of bearing 
vestigial, peg-like structures (Ichinose, 19626; Eichlin & Cunningham, 1978). All species of Acanthoplu- 
sia, except A. agnata and A. latistigma, have a similar, very finely reticulate, brown forewing pattern. 

Ctenoplusia (s.str.) [4(3), 60(2), 78(2), 86(0)]. As mentioned above (Character analysis, character 60), 
Ctenoplusia (s.str.), as currently constructed, is probably a polyphyletic assemblage of those species with 
blade-like saccular setae (and some that lack them) that are not members of Acanthoplusia or Plusiotricha. 
The last genus shares a similar form of basitarsal comb not found in Acanthoplusia (see character 60, 
Character analysis), while the conspicuous pink forewing spot appears to be a potential synapomorphy for 
C. limbirena (Fig. 54) and several other species (see character 4, Character analysis). The distribution of 
dorso-lateral tufts that are enclosed in a pleural fold (78) needs to be checked across a wider range of 
species. 

Of the 50 or so species of Ctenoplusia (s.str.), the life histories of only two appear to have been 
elucidated. C. limbirena, like many plusiines, is polyphagous, feeding on Althea (Malvaceae), Becium, 



148 IAN J. KITCHING 

Salvia (both Lamiaceae), Primula (Primulaceae), Verbascum (Scrophulariaceae), Solatium and Nicotiana 
(both Solanaceae) (Pinhey, 1975). In contrast, C. albostriata (which is possibly not congeneric with C. 
limbirena) is apparently restricted to members of the Asteraceae (Gardner, 1947; Robinson, 1975; 
Miyata, 1983; Holloway, 1985), although there are records for Calystegia (Convolvulaceae) (Miyata, 1983) 
and Symphytum (Boraginaceae) (Holloway, 1985). Further support for a close relationship between C. 
albostriata and Trichoplusia vittata and T. oxygramma (see 60, Character analysis) is provided by Pinhey's 
(1975) record of Commidendron (Asteraceae) as the foodplant of T. vittata (as transfixa) and Crumb's 
(1956) records of Aster, Solidago, Erigeron (Asteraceae) and Nicotiana as foodplants of T. oxygramma. In 
C. albostriata, a form is known (f. innotata) in which the modified Y-mark is missing. The genetics of this 
and the typical form (f. albostriata) was investigated by Ichinose etal. (1967). The two forms were found to 
obey Mendel's laws of heredity, with f . albostriata being totally dominant to f . innotata. 

Clade 17 [61(1), 83(4), 87(0), 138(0)]. Plusiopalpa, Stigmoplusia and Argyrogramma are united as clade 17 
on the basis of character 61(1), a suppressed inner SV row of setae on the male foretarsi. This state is, 
however, convergently derived in Plusiotricha. 

Plusiopalpa [12(1), 41(1), 84(0), 86(0), 91(1), 128(1), 201(1), 213(2)]. Plusiopalpa is a compact genus of 
five species that have an unmistakable general appearance. In addition to large size and a somewhat 
distinctive colour pattern, their most obvious feature is that which has given the genus its name, the very 
long labial palps. This is due to the extreme dorso-apical elongation of the third segment (41, see Character 
analysis) . In addition there are numerous generic autapomorphies of the genitalia of both sexes including a 
scimitar-shaped valve, vinculum arms with a marked dorsal keel and medial constriction, and a distinctive 
corpus bursae in which the ductus seminalis arises from a small apex produced laterally about halfway 
down the corpus bursae, near the point of entry of the ductus bursae. 

Plusiopalpa occurs throughout the Old World tropics. P. thaumasia Dufay and P. hildebrandti 
(Saalmuller) are endemic to Madagascar, while P. shisa appears to be restricted to Taiwan. P. adrasta 
ranges throughout the Oriental tropics to New Guinea, and P. dichora (Fig. 58) occurs in Africa. 
Hampson's (1913) synonymy of P. adrasta and P. dichora, followed by Gaede (1940) and Pinhey (1975), is 
incorrect (Dufay, 1970a), while the separation of P. shisa from P. adrasta ought to be confirmed by 
genitalic dissection. At present, the only diagnostic characters are minor wing pattern elements. Hampson 
(1913) also included a sixth species in Plusiopalpa, P. camptogamma (Hampson). Dufay (1970a) disagreed 
with this placement but did not suggest a better one. Examination of the genitalia of the female holotype 
showed that it does not possess the characteristic form of the corpus bursae referred to above that is found 
in other plusiopalpas, but has a simple corpus bursae instead. Thus P. camptogamma should be removed 
from Plusiopalpa. Dufay (unpublished combination in BMNH collection) has proposed that the species is 
properly placed in Ctenoplusia, and the possession by the male of a quadrisetose basitarsal comb (see 
Character analysis) supports this move. However, examination of the male genitalia is required to confirm 
this placement as correct. Only P. adrasta has ever been reared, from Mikania, an asteraceous plant 
introduced into Peninsular Malaysia (Holloway, 1985). 

Clade 18 [60(3), 61(2), 79(3), 85(1), 90(1), 92(2), 93(3), 94(3), 130(0), 162(1)]. Although outwardly 
different in general appearance, Argyrogramma (s.str.) and Stigmoplusia share a remarkable number of 
synapomorphic features, all of which are probably secondary sexual characteristics of the males. These are: 
a basitarsal comb composed of multiple rows of setae (60) ; a further suppression of the foretarsal S V spines 
(61); dorso-lateral hair tufts with discrete edges on segment A6 only (79, but see Agrapha and Stigmoplu- 
sia); differentiated sternal hair pencil insertions (85); an anterior flange on the lateral arm of St8 at the 
point where they curve dorsally over the hair pencils (90); lateral arms of St8 with the sub-basal lobe 
modified into a blunt projection (92); dorsal arms of St8 that run almost transversely (93); and apical scales 
on the dorsal arms of St8 modified into a semicircular, projecting plate (94). 

Argyrogramma (s.str.) [61(3), 78(3), 88(1), 98(1), 105(1), 108(0), 121(2)]. The members of the genus 
Argyrogramma (s.str.) are most easily recognized by a feature of the male genitalia - the vinculum arms 
bear broad dorsal and ventral flaps that are folded horizontally and partially fused together to give a stocky, 
rectangular saccus that is markedly truncate anteriorly (e.g. A. signata; Dufay, 1970a: fig. 96). In this 
study, two autapomorphies were found for A. verruca, characters 61 (completely suppressed male 
foretarsal SV spines; short, stout V spines) and 78 (dorso-lateral hair tufts on A7 enclosed in a membranous 
fold). Two further character states (88, 98) were shared only with Agrapha (q.v.) and one only with 
Abrostola (121). Their occurrence in the other members of the genus needs confirmation. 

Argyrogramma (s.str.) currently contains 12 species. However, when first described (Chou & Lu, 1978; 
1979a), five were placed in Argyrogramma in the broad sense (i.e. sensu Ichinose, 1973). These species are 



SYSTEMATICS OF THE PLUSIINAE 149 

more likely to prove to be ctenoplusias or thysanoplusias and I will not consider them further. A. 
circumscripta does not agree with the type species, A. verruca, either. However, Kostrowicki (1961), who 
transferred circumscripta to Argyrogramma, had a very peculiar concept for the genus and this species also 
probably belongs elsewhere. Similarly, A. aenofusa was transferred to Argyrogramma (s.l.) by Chou & Lu 
(1978). However, this species does share a potential apomorphy with A. verruca, namely a basitarsal comb 
composed of multiple rows of setae (60:3, see Character analysis). Thus, for the moment, I will leave A. 
aenofusa in Argyrogramma (s.str.), despite numerous other differences (e.g. highly modified, spatulate 
saccular setae). 

The core of Argyrogramma (s.str.) can thus be considered to consist of five species. Of these, A. 
hainanensis Chou & Lu is almost certainly synonymous with A. signata, a widespread species in the Old 
World tropics, found from the Canary Islands, through to Tonga and the Cook Islands (Dufay, 1970a; 
Holloway, 1985). A. subaerea Dufay is recorded from widely scattered localities in tropical Africa from the 
Ivory Coast to Uganda and SW. Angola (Dufay, 1972), while A. verruca (Fig. 59) and A. basigera occur 
throughout North America and Mexico, with the former extending down to Peru and Argentina (Eichlin & 
Cunningham, 1978), even reaching the Galapagos Islands (Hayes, 1975). 

The larvae of both A. verruca and A. basigera have been described by Eichlin & Cunningham (1978). 
The former is polyphagous while the larval hostplant of the latter has not been recorded. Although the 
immature stages of A. signata have not been described, Holloway (1985) stated that the species has been 
reared on Eucalyptus in New Guinea. Given that most plusiines feed on herbaceous plants, this record 
requires confirmation as a natural hostplant. 

Stigmoplusia [11(1), 79(2), 83(3), 97(1), 111(1), 112(2), 144(1), 214(0)]. Three synapomorphies were 
found for Stigmoplusia chalcoides in the present data set: discrete edges to the dorso-lateral hair tufts on 
segment A5 only (79) and a vinculum produced as a long spine ( 1 1 1 ; similar to the condition found in clade 
8, q.v.), bearing two sharp, subdorsal points (112). There are also several autapomorphies not contained in 
the data set, including a highly differentiated sternal hair pencil comprising several types of scales (Fig. 
270); broad, subventral flaps on the uncus (Figs 280, 281); uncus apically rounded-triangular in cross- 
section; and strong, asymmetrical digitations on the ventral edge of the valve, distal to the apex of the 
sacculus, which terminates in a stout, curved spine (Fig. 353). 

Seven species are currently placed in Stigmoplusia, all described by Dufay (1970a; 1972; 1975): acalypta, 
allocota, antsalova, chalcoides (Fig. 60), epistilba, megista and paraplesia. All except 5. acalypta and S. 
megista possess uncal flaps. In addition to 5. chalcoides, valve digitations are present in S. allocota, S. 
acalypta and S. megista, but occur along the entire ventral edge of the valve, each digitation bearing a 
single, short, apical seta. 5. epistilba and S. paraplesia have simple valves, while those of 5. antsalova are 
very peculiar (see Dufay, 1970a: fig. 109); the apex is enlarged with a basally-oriented, rounded lobe that 
bears an apical membranous sac. As far as can be ascertained, all species have vinculum arms with 
subdorsal, sharp points. Stigmoplusia thus presents quite a diverse array of male genitalic structure. The 
females of only three species are known (S. chalcoides, S. allocota, S. acalypta) and all agree in the apex of 
the corpus bursa being ornamented internally with a conspicuous, sclerotized, reticulate pattern of ridges. 

Nothing is known about the biology of any species of Stigmoplusia. The genus is found in the tropical 
forest belt of Africa from Liberia to Uganda and Zimbabwe. S. antsalova, with the unusual male genitalia, 
is endemic to west Madagascar. 

Clade 19: Plusiini [117(1), 121(1), 127(1)]. The members of the primarily Holarctic tribe Plusiini are united 
by the synapomorphic possession of a broad flange on the pleurite (117). This structure is, however, 
subsequently lost in the highly autapomorphic Antoculeora. A juxta with an apical notch (or subsequent 
derivation) (127) is also characteristic of this tribe, although it is lost in Allagrapha and convergently 
derived in Omorphina and Chrysodeixis . 

The larvae of the Plusiini are characterized by the total loss of the prolegs on segments A3 and A4 (but 
these are also convergently lost in Acanthoplusia agnata, see above). There also appears to be a greater 
degree of larval oligophagy in the Plusiini, but this may be due solely to a lack of information for many 
(particularly non-pest) members of the Argyrogrammatini. 

Clade 20: Euchalciina [39(1), 83(0), 101(1), 178(2)]. The genera of the plusiine subtribe Euchalciina are 
characterized by two good synapomorphies: the pleural pouch is small and concealed within the tympanal 
hood (178) and all have a distinctive outline to the tegumen in lateral view (101). In addition, the sternal 
hair pencils are invariably reduced to a mere sparse tuft of scales (83), although this reversal to the 
plesiomorphic condition has also taken place in other genera (Pseudoplusia, Eutheiaplusia, Syngrapha). 
Finally, labial palps in which the ventral scales form a neat ridge (39) are also characteristic of the 



150 IAN J. KITCHING 

Euchalciina, but the condition appears to have been lost in Euchalcia and Adeva (clade 25). Most of the 
euchalciines feed as larvae on Ranunculaceae, which may prove to be a further synapomorphy for the 
subtribe. 

Eosphoropteryx [175(1), 213(1)]. Eosphoropteryx is a monobasic genus, comprising E. thyatiroides (Fig. 
61). It has a number of genitalic autapomorphies, including a keel-shaped valve with a produced apex; an 
unsclerotized , sub-basal auricular cornutus on the vesica ( 175 ; similar cornuti occur in Macdunnoughia and 
Loboplusia, but are heavily sclerotized in these genera); a flattened, laterally curved, heavily sclerotized 
antrum (Fig. 452); and granular ridges on the apex of the corpus bursae (213). 

E. thyatiroides is a pinkish-grey plusiine, found across the northern United States and southern Canada, 
from Nova Scotia and Virginia to British Columbia and Oregon (Eichlin & Cunningham, 1978). The larva 
is reported to feed on Thalictrum (Ranunculaceae) (Forbes, 1954) but no other details of the species' 
biology are known. 

Clade 21 [109(1), 147(1), 196(1)]. The remaining genera of the Euchalciina, excluding Eosphoropteryx, are 
united primarily by the synapomorphic shape of the valve (147). The simple, oval valve occurs only in clade 
21, with the exception of a convergent occurrence in Omorphina (q.v.). As noted above (Character 
analysis), the valve of Euchalcia variabilis is more quadrate than oval, but the majority of euchalcias show 
the oval valve form. An S-shaped vinculum apex in lateral view ( 109) is weakly expressed in this clade and 
also occurs in clade 39 (Plusiina part) and in Cucullia. A broad, shallow, open, bowl-like antrum (196) is 
also characteristic of all members of clade 21 but is highly homoplasious, occurring additionally in Agrapha 
and clade 8 within the Plusiinae and in Diloba, Cucullia and Chalcopasta in the outgroups. 

Polychrysia [5(1), 41(1), 50(1), 65(2), 165(1)]. One unique and unreversed apomorphy for Polychrysia was 
found in the present data set; the presence of a few, fine SD and D setae on the tarsi (65). It is unknown if 
this character state is present in the other members of the genus. Polychrysia also possesses scattered black 
forewing scales (5), an elongate labial palp segment 3 (41) and erect tegulae (50), all of which occur 
(homoplasiously) only in the Euchalciina (with the exception of the first, which is also found elsewhere in 
the Plusiini). The subapical cornutus borne on a lateral diverticulum (165) may represent another 
apomorphy for the genus. The superficially similar condition seen in Diachrysia is almost certainly not 
homologous. 

Polychrysia currently contains six species. The most widespread is P. moneta (Fig. 62), known in Britain 
as the Golden Plusia. It occurs across Canada from Ontario to Alberta, and north to Alaska, but has yet to 
be recorded from British Columbia or east of Lake Huron (Eichlin & Cunningham, 1978). Widely 
distributed in Europe, P. moneta is replaced east of the Urals by P. esmeralda (Oberthiir). This species 
differs from P. moneta in the silver-grey ground colour to the forewing, rather than a golden-yellow, and 
has only recently been reinstated as a good species (Kostrowicki, 1961; Chou & Lu, 1979a). However, 
there is some confusion (compare the figures of the male genitalia in the above two publications) and the 
separation of P. esmeralda from P. moneta is still open to question. 

P. sica, P. aurata (Staudinger) and P. splendida (Butler) are all eastern Palaearctic (China, Japan, 
Siberia) (Kostrowicki, 1961), while P. morigera (Edwards) is restricted to the foothills of the Rocky 
Mountains of the U.S.A., both the eastern and western slopes. 'Polychrysia' aureus (Bang-Haas) 
(Kostrowicki, 1961) is not a plusiine but is correctly assigned to the amphipyrine genus Chrysonicara 
Draudt, near Nicara Moore (Draudt, 1937). 

The life history of P. moneta in Europe is well known (Bretherton et ai, 1983). The larvae are most 
frequently recorded feeding on members of the Ranunculaceae; e.g. Aconitum, Delphinium (South, 1961; 
Skinner, 1985) and Trollius (Beck, 1960; Kostrowicki, 1961). However, the last author also gives Cucumis 
(Cucurbitaceae) and Helianthus (Asteraceae), while South (1961) adds Arctium (Asteraceae). P. splendi- 
da appears to be restricted to Aconitum (Miyata, 1983), while P. morigera is recorded only from 
Delphinium sp. (Eichlin & Cunningham, 1978). Pupation is in a characteristic strong, oval cocoon of 
yellow silk spun between leaves of the foodplant (Bretherton et al. , 1983). 

Clade 22 [3(0), 154(1), 183(1)]. The members of clade 22 are primarily characterized by short anal papillae 
(183). However, as noted above (Character analysis), Euchalcia variabilis exhibits the plesiomorphic 
condition of approximately square papillae and this was interpreted as an early stage in a transform- 
ation series culminating in the blade-like papillae of species such as E. augusta. Transtillae that are fused 
above the aedeagus into a broad, blunt point (154) also characterize clade 22, with the exception of the 
included clade 26. Further investigation may show this character state to be a synapomorphy of clades 23 
and 25. 



SYSTEMATICS OF THE PLUSIINAE 151 

Clade 23 [42(1), 132(1), 163(1)]. Two synapomorphies unite Panchrysia and Chrysanympha into a 
monophyletic group: fused saccular flanges modified into a rounded lobe between the valves (132) and an 
apically swollen subapical cornutus (163). However, the latter appears to be present in Panchrysia only in 
P. deaurata, P. ornata (Bremer) (Kostrowicki, 1961; Chou & Lu, 1978) and P. mishanensis Chou & Lu 
(Chou & Lu, 1979a), while the former may be of more general occurrence, being distinguishable in figures 
of the genitalia of P. deaurata, P. ornata and P. dives but not P. v-argenteum (Esper) (Kostrowicki, 1961) 
or P. mishanensis (Chou & Lu, 1979a). The sister-group relationship of Panchrysia and Chrysanympha 
therefore remains equivocal. 

Chrysanympha [5(1), 41(1), 107(1)]. Chrysanympha contains a single species, C. formosa (Fig. 63), of 
unique general appearance within the Plusiinae. The forewing has a strong, brown, evenly-curved 
subterminal line and the pinkish grey colour of the Y-mark is extended costally and basally to give a 
superficially similar pattern at the wing base to Abrostola (into which genus C. formosa was incorrectly 
placed by Hampson, 1913). Chrysanympha also bears a dorsal sclerotized bar on the anal tube (107), 
similar to that seen in Lophoplusia and the Abrostolini, and is another euchalciine to display scattered 
black scales on the forewing (5). 

C. formosa appears to be a scarce species confined to the north-eastern U.S.A. and south-eastern 
Canada (Eichlin & Cunningham, 1978). Its life history has been briefly described by Kearfott (1904) and 
the larval foodplants are given as Gaylussacia and Vaccinium (Ericaceae) (Tietz, 1972), unusual for the 
Euchalciina. 

Panchrysia [19(1), 65(3), 209( 1 )] . Panchrysia deaurata is the only euchalciine to my knowledge in which the 
ductus seminalis arises from the fundus of the corpus bursae (209). This may prove to be an autapomorphy 
for the species, as the ductus seminalis has the more usual origin from the apex of the corpus bursae in P. 
ornata (Chou & Lu, 1978: fig. 23). Like Polychrysia, Panchrysia deaurata has SD and D tarsal setae (65) 
but in the present genus, they are numerous and long, much as in the distantly related Lophoplusia. The 
presence of this character state in the other members of the genus needs confirmation. 

Five species currently comprise Panchrysia. The type species, P. deaurata (Fig. 64), is found in steppe 
and semi-arid areas of Europe and Central Asia from Spain to the Altai. It is a golden-yellow moth, 
superficially(?) similar to Polychrysia aurata, and thus unlike the other members of its genus in general 
appearance. The other four species all bear a silver Y-mark and other metallic spots on the forewings. P. 
v-argenteum (Fig. 65) is a pinkish brown moth restricted to the European Alps and the high mountains of 
south-east Europe (e.g. Yugoslavia, Greece), while P. ornata is very similar but pale grey-brown. The 
latter species occurs in the eastern Palaearctic steppes (Kostrowicki, 1961), and thus stands in relation to P. 
v-argenteum, in terms of both distribution and colour, much as Polychrysia esmeralda stands in relation to 
Polychrysia moneta. Panchrysia mishanensis is very similar to P. ornata (Chou & Lu, 1979a) and 
presumably originates from Mi-Shan in far north-eastern China, but this is not certain. The fifth species, P. 
dives (Fig. 66), is similar to P. v-argenteum but the forewing is chocolate-brown, has more extensive silver 
markings, and the hindwing is yellow with a brown margin. It is largely sympatric with P. ornata, but 
extends further east into southern Kamtchatka and is absent from the Korean peninsula and most of China 
(Kostrowicki, 1961). 

Little is known of the biology of Panchrysia. The larval foodplants of P. deaurata are recorded as 
numerous species of Thalictrum (Ranunculaceae), while P. v-argenteum feeds on Isopyrum (Ranuncu- 
laceae), as well as Thalictrum foetida (Kostrowicki, 1961). 

Clade 24 [6(0), 144(1), 168(1), 205(1)]. The five genera comprising clade 24 are united by the synapomor- 
phic possession of a vesica that is internally granular basally (168). 

Clade 25 [4(4), 39(0), 58(0), 164(1)]. Euchalcia and Adeva are considered sister-taxa on the basis of two 
synapomorphies. In both genera, the terminal line on the forewing is white not pink (4) and the subapical 
cornutus on the vesica arises from a circular, concave base-plate (164). These two character states also 
occur in all other Euchalcia species represented in the BMNH collection or figured by Dufay (1968), with 
the following exceptions. The white terminal line is absent in E. inconspicua (Graeser) (Fig. 69), E. 
cashmirensis Moore and E. dorsiflava, and is reduced in several other species. The circular base-plate of 
the subapical cornutus is absent only in E. inconspicua, where the subapical cornutus is replaced by a series 
of cornuti, and E. tancrei (Staudinger). Dufay (1968) considered Euchalcia to be a very homogeneous 
genus, in terms of general facies and male and female genitalia, providing E. inconspicua, E. tancrei and 
'Plusia' hampsoni Leech (Fig. 68) were excluded (all were first transferred to Euchalcia by Kostrowicki, 
1961, the last as its synonym, 'Phytometra' lenzeni Draudt [Dufay, 1965]). Euchalcia inconspicua has 



152 IAN J. KITCHING 

recently been placed in its own genus, Pseudochalcia , by Kljutshko (1984). I have examined the male 
genitalia of hampsoni (BM noctuid slide no. 11521) and agree with Dufay (1968) that this species does not 
belong in Euchalcia. I am unable at present, however, to suggest where 'P.' hampsoni ought to be placed. I 
superficially examined specimens of Euchalcia tancrei, together with the drawing of the male genitalia in 
Kostrowicki (1961), and I concur with Dufay that this species, also, does not belong in Euchalcia. Although 
it is unsatisfactory, until such time as a more comprehensive study is performed, both hampsoni and tancrei 
should be placed in limbo (i.e. 'Plusia' s.l.). This action, together with the removal of inconspicua to 
Pseudochalcia (see also Consensus classification), almost certainly leaves Euchalcia (+ Adeva) 
monophyletic. 

Euchalcia [183(0), 201(0)]. The genus Euchalcia comprises around 30 species. They are found almost 
exclusively in the steppe and semi-arid regions of the Palaearctic, the exceptions being E. cashmirensis , E. 
serraticornis Dufay and E. xanthoides Dufay, which are subtropical-Himalayan species (Dufay, 1968). 
Euchalcia gerda and E. serraticornis have serrate male antennae (the females are unknown for both 
species), and are thus somewhat unusual (Dufay, 1968). 

The biology of only seven species is known (Dufay, 1968). E. variabilis (Fig. 67) and E. bellieri (Kirby) 
are reported to feed as larvae on the usual ranunculaceous foodplants of the Euchalciina, Aconitum and 
Thalictrum. However, one group of euchalcias has undergone a foodplant switch to members of the 
Boraginaceae: Alkana (E. chlorocharis (Dufay)), Cynoglossum (E. cuprea (Esper) and E. italica (Staudin- 
ger)), Lycopsis (E. consona (F.)), Pulmonaria (E. consona and E. cuprea), Rindera (E. siderifera 
(Eversmann)), Nonea (as Nonnea; E. consona) and Symphytum (E. cuprea) (Beck, 1960; Kostrowicki, 
1961; Dufay, 1968). In addition, E. cuprea is recorded from Pterotheca (Asteraceae) and, most unusually, 
Arum (Araceae) (Kostrowicki, 1961). 

A comprehensive revision of the genus has been provided by Dufay (1968), including detailed accounts 
of the morphology, distribution and affinities of 29 species of Euchalcia and separate keys to species using 
general habitus, male and female genitalia. 

Adeva [5(1), 59(0)]. Adeva is yet another euchalciine to have scattered black scales on the forewing, and in 
this character state, the sole included species, A. albavitta, differs from all species of Euchalcia that I have 
examined. 

Adeva was erected by McDunnough (1944) to accommodate A. albavitta (Fig. 71). The genus was 
considered synonymous with Euchalcia by Kostrowicki (1961), a decision endorsed by Dufay (1968). A. 
albavitta is restricted to the south-western U.S.A., where it occurs in the states of Oregon, California, 
Arizona and Nevada (Eichlin & Cunningham, 1978). There are two colour forms, a pale brownish form 
found in the coastal regions of California, and a smoky grey desert form, originally described in the genus 
Behrensia Grote (McDunnough, 1944). Nothing is known about the biology of A. albavitta. However, 
given the dichtomy in Euchalcia of larval foodplants between the Ranunculaceae and the Boraginaceae, 
knowledge of that of Adeva might prove useful in resolving the position of this species with regard to the 30 
or so euchalcias. 

Clade 26 [144(2), 154(0), 171(1), 186(1), 194(2)]. Plusidia, Lamprotes and Pseudeva form a monophyletic 
group on the basis of three synapomorphies: the presence of an isolated, dorsal patch of cornuti on the 
vesica (171); short, thick membrane preventing easy protraction of the anal papillae (186); and the 
presence of several setae along the inner, ventral edge of the antrum (194). This last character state is 
further transformed in Lamprotes (q.v.) and a superficially similar but non-homologous condition occurs in 
Diloba. 

Plusidia [58(0)]. Plusidia currently contains three species. However, P. imperatrix and P. chinghaiensis 
Chou & Lu show considerable similarities in wing pattern to Polychrysia splendida and examination of the 
genitalia may show that they are properly placed in Polychrysia. 

This action would render Plusidia monotypic, containing only P. cheiranthi (Fig. 34). At one time, the 
eastern Palaearctic populations were considered to be a separate species, P. separanda Warren (Inoue & 
Sugi, 1958-61). More recently, it has been relegated to subspecies of P. cheiranthi (Kostrowicki, 1961) and 
finally considered to be fully synonymous (Sugi, 1982). 

Western populations of P. cheiranthi are unmistakable in appearance. The forewings are pinkish grey, 
with a chocolate-brown, comma-shaped apical spot and a similar brown spot at the base. However, eastern 
examples have a much more 'washed-out' appearance with the apical spot barely distinguishable and the 
basal spot often somewhat faded also. The larva was briefly described by Kirby (1889), who gave the 
foodplants as Thalictrum and Aquilegia (Ranunculaceae). 



SYSTEMATICS OF THE PLUSIINAE 153 

Clade 27 [6(1), 16(1), 19(1), 41(1), 42(1), 205(0)]. There are no good synapomorphies to support the 
sister-group relationship between Lamprotes and Pseudeva, which must thus be considered tentative at 
best. Nevertheless, this relationship has been proposed previously by Kostrowicki (1961), who stated that 
Lamprotes was 'in facies and genitalia much like the species of the Nearctic genus Pseudeva' (see also 
Pseudeva below). 

Lamprotes [3(1), 144(1), 162(0), 191(1), 194(1), 201(1)]. Lamprotes can be separated from the other two 
genera of clade 26 by the restriction of the ventral antrum setae to a subventral or lateral position (194). In 
addition, most Lamprotes specimens bear a small, golden, C-shaped mark below the forewing cell (3). This 
marking was coded in the data as homologous with the Y-mark of other plusiines. However, the C-mark of 
Lamprotes occupies a slightly more basal position (entirely below vein CuA 2 rather than straddling it) and 
this suggests that it represents a novel pattern element that is autapomorphic for Lamprotes. 

Lamprotes has a very similar disjunct distribution to Plusidia, with western and eastern Palaearctic 
populations. However, in contrast to that genus, the eastern population of Lamprotes, L. mikadina Butler, 
has long been treated as a good species (Hampson, 1913; Inoue & Sugi, 1958-61; Chou& Lu, 1979a; Sugi, 
1982). Occasionally, L. mikadina has been treated as a subspecies of L. c-aureum (Kostrowicki, 1961) (Fig. 
35), or even fully synonymized (implicitly in Forster & Wohlfahrt, 1971). 

The larva of L. c-aureum has been briefly described by Kirby (1889) and that of L. mikadina in more 
detail by Ichinose (19626). Both species are reported to feed on Thalictrum and Aquilegia. 

The generic name Lamprotes is unusual insofar as the identity of the author is unknown, only his or her 
initials accompanying the first reference to the name (R.L., 1817). According to W. H. T. Tarns (Nye, 
1970; unpublished notes), the identity of R.L. was investigated by C. D. Sherborn. However, he reached 
no firm conclusion. C. Boursin has attributed authorship to Reichenbach of Leipzig, but it could also be 
Rudolph Leuckhart (who published under his own name 1836-1859) or Rudolph Ludwig, suggested by E. 
L. Todd (Nye, 1970; unpublished notes). Nye suggested that continuing to use R.L. was the most prudent 
course. 

Pseudeva [5(1), 50(1)]. Pseudeva contains only two species, P. purpurigera (Fig. 36) and P. palligera. Both 
species, like Polychrysia, have erect tegulae (50). However, this character state is difficult to score in any 
specimen that is little less than perfect and may yet be shown to occur in other euchalciine genera. 

Both species of Pseudeva are endemic to the Holarctic region. P. palligera occurs only in the 
south-western U.S.A. , while P. purpurigera is widespread in southern Canada and the northern two-thirds 
of the U.S.A., east of the Rocky Mountains, but is noticeably absent from the central band of states from 
North Dakota to Texas. 

The larva of P. purpurigera has been described by Dyar (1896), who gave Thalictrum as the foodplant. 
He also stated that the larva was 'green and white [and] curiously hunched up'. This description is very 
similar to the coloration and posture of the larva of Lamprotes mikadina (Ichinose, 19626: pi. 18, fig. 4; 
Mutuura etal. , 1965: pi. 37, fig. 113) and these features may represent good synapomorphies for Lamprotes 
and Pseudeva. Further examination of the larvae of Pseudeva is thus important in confirming the 
sister-group relationship of these two genera. 

Clade 28 [116(1)]. The subtribes Plusiina and Autoplusiina are interpreted as sister-taxa solely on the basis 
of a strongly curved pleurite ( 1 16) . As noted earlier, such a structure also occurs in Anadevidia and thus the 
monophyly of clade 28 can only be considered to be weakly supported. Eichlin & Cunningham (1978) 
treated the Plusiina and (paraphyletic) Euchalciina as sister-taxa on the basis of a shared fusion of the 
pinacula of setae SV1 and SV2 on segment A2 of the larva. However, this character state also occurs in 
several non-Nearctic autoplusiine genera (Macdunnoughia, Sclerogenia and Erythroplusia), as well as in 
the argyrogrammatines Thysanoplusia and Anadevidia (Ichinose, 19626), and is thus of dubious value. 

Clade 29: Autoplusiina [18(0), 213(2)]. The members of the Autoplusiina are only weakly united as a 
monophyletic group on the basis of sclerotized ridges at the entry point of the ductus bursae into the corpus 
bursae and on the apex of the corpus bursae (213). However, these ridges must have been secondarily lost 
in Erythroplusia. In addition, such ridges occur in Omorphina and Plusiopalpa within the Plusiinae, and in 
Diloba and Chalcopasta of the outgroups. Further study is required into this character and, given the 
reservations regarding the homologies of other character states important in the classification of this 
subtribe (e.g. see Characters, 142: 1 & 2), the Autoplusiina may subsequently be shown to be paraphyletic 
or worse. 

Rachiplusia [14(2), 59(0), 63(1), 64(2), 110(1), 114(1), 139(1), 148(0). 154(1), 191(1), 198(3), 206(1), 



154 IAN J. KITCHING 

216(1)]. Rachiplusia is perhaps the most problematical autoplusiine genus, in that it shares possible 
synapomorphies with a number of subgroups of the subtribe. Rachiplusia and 'Syngrapha' gammoides are 
the only autoplusiines that are sexually dimorphic (216), while a bisetose female frenulum (14) and spined 
hind tibiae (64) further corroborate a relationship between Autoplusia and Rachiplusia. Rachiplusia also 
shares character states with larger clades that include Autoplusia: increased sclerotization of the LPV and 
T8 ventrally (198: clade35);lackof aharpe(140: clade34); and a protected ostium bursae (191: clade36). 
This last character is also shared, however, with the members of clade 32, as is a clavus base that is obscured 
by a saccular overfold (139). Finally, Rachiplusia and Macdunnoughia s.l. (clade 33) both have a medially 
expanded section of the ductus bursae with dorsally recurved edges (206). It is thus unlikely that 
Rachiplusia will prove not to be related to some or all of the autoplusiines of clade 30. However, the genus 
lacks many structures found in this clade, most notably the apomorphic states of characters 95, 141 , 189 and 
193 (see below) . A more extensive survey of the Autoplusiina (and maybe the Plusiina, seeAnagrapha and 
Syngrapha below) is necessary to resolve these ambiguities. 

Rachiplusia comprises three species. Two, R. nu (Fig. 72) and R. ou. are very similar, both in general 
appearance and in genitalia. R. ou occurs from the U.S.A. south to Venezuela (including Jamaica and the 
Bermuda Islands), while/?, nu is found in Bolivia, Chile, Brazil and Argentina. The immature stages of R. 
nu have been described in detail by Angulo & Weigert (1974), and the larvae of R. ou by Crumb (1956). 
Both species are highly polyphagous (Angulo & Weigert, 1975; Eichlin & Cunningham, 1978) but only R. 
nu appears to be of economic importance as an agricultural pest (Angulo & Weigert, 1975). 

The third species, R. virgula, has long been considered a member of the genus Caloplusia (e.g. 
Hampson, 1913; Kostrowicki, 1961), because of its reduced eyes, yellow hind wings and fully spined tibiae. 
Following examination of the genitalia, however, Angulo (1978) transferred virgula to Rachiplusia. There 
can be little doubt that this reflects the relationships of the species more precisely. Both R. virgula and R. 
nu (the second species figured by Angulo, 1978) have a ventrally bulged valve (Fig. 360); long, thin clavi 
(Fig. 360); a broad, truncate saccus (Fig. 302); and a similar arrangement of cornuti on the vesica. The 
immature stages of R. virgula have been described by Izquierdo (1895), who gave the larval foodplant as 
Madia sativa (Asteraceae). R. virgula is endemic to the Chilean Andes. 

It does not appear to have been noted before that two of the species of Rachiplusia are sexually 
dimorphic. The males are more brightly and contrastingly coloured than the females, which are dull in 
comparison. In addition, the males have a predominantly grey forewing ground colour, while the females 
are brown. A similar but more striking dimorphism occurs in 'Syngrapha' gammoides (Figs 82, 83). In 
contrast, R. virgula appears to be monomorphic, although I have not seen a sufficiently long series of this 
species to be certain. 

Clade 30 [83(2), 95(1), 141(1), 162(0), 189(1), 193(1)]. The remaining members of the Autoplusiina, 
excluding Rachiplusia, are united by a single unique and unreversed synapomorphy, the presence of short, 
upright scales on the antero-ventral margin of the antrum (193). In addition, a broad, triangular saccular 
spine (141) occurs in many members of the subtribe, although some extreme modifications may eventually 
prove to be incorrectly homologized (e.g. the quadrate lobe of Erythroplusia and the lance-like projection 
of Antoculeora) . Fine, hair-like scales ventrally between the lateral arms of St8 (95) occur in all the member 
genera of clade 30 except Sclerogenia, while a T8 that is cut back ventral to the anterior apophysis (189) is 
present in all but Antoculeora, where the extreme autapomorphic condition of the male genitalia make it 
difficult to score the character. However, the apomorphic state of character 189 also occurs in Syngrapha 
(s.l.) and Anagrapha (see below). Again, a more extensive investigation is necessary, particularly of the 
male genitalia, in order to confirm several of the homologies hypothesised in this study (especially those of 
characters 141 and 142). 

Erythroplusia [5(1), 6(1), 18(1), 104(1), 108(1), 145(1), 155(1), 158(2), 209(1), 213(0)]. Like Rachiplusia, 
the relationships of Erythroplusia are somewhat obscure. It also possesses character states that associate it 
with several different members of the Autoplusiina. Convergent upon the condition found in Antoculeora, 
the tegumen arms of Erythroplusia, in dorsal view, diverge at about 180 degrees, rather than forming the 
more usual inverted V-shape (104). Erythroplusia bears its clavus on a long, flattened peninsula ( 145) , as in j 
Antoculeora and Macdunnoughia (s.l.). As in the latter genus (s.str.), the ventral sclerotization on the j 
aedeagus of Erythroplusia is restricted to a very thin, ventral band, convergent upon the condition seen in 
the Argyrogrammatini. Finally, the form of the transtillae (155) is almost identical to that seen in 
Sclerogenia. 

In addition to the above states, the two constituent species of Erythroplusia show several other 
remarkable genitalic autapomorphies. The valves are asymmetrical (Fig. 378). The sacculus of the left 
valve bears a distally directed, curved process, while that of the right valve bears a subapically recurved. 



SYSTEMATICS OF THE PLUSIINAE 155 

heavily-sclerotized ridge and no process. The corpora bursae of the females are highly simplified relative to 
those seen in other autoplusiines. That of E. rutulifrons (Fig. 453) is a simple, cylindrical sac, somewhat 
swollen where it recurves posteriorly through about 180 degrees. There is no obvious differentiation into 
ductus and corpus bursae. The corpus bursae of E. pyropia is more swollen and tapers distally. In both, it is 
extremely difficult to ascribe the terms apex and fundus with any degree of certainty (see also Characters: 
209). 

Erythroplasia was originally proposed as a subgenus of Autographa (Ichinose, 1962a; b). Subsequently, 
it was elevated to full generic rank (Ichinose, 1973) and included Antoculeora (q.v.) as a subgenus. The 
precise relationships of Erythroplasia are still equivocal. There are some larval characters that suggest a 
relationship with the Argyrogrammatini (a slender seta SD1 on segment A9 was used by Kljutshko, 1985a, 
to associate the two taxa as sister-groups), while the chaetotaxy is identical in Erythroplasia and 
Autographa (which included Sclerogenia; Ichinose, 1973). 

The larvae of both species have been described by Ichinose (19626). No foodplants were given for E. 
rutulifrons but that of E. pyropia was stated to be Artemisia (Asteraceae). Miyata (1983) gave Arctium 
(Asteraceae) and Oenanthe (Apiaceae) for both species. Erythroplasia is primarily north-eastern 
Palaearctic in distribution, with the two species being broadly sympatric over much of their ranges. 
However, E. pyropia extends further west than E. rutulifrons (Fig. 73), occurring throughout the southern 
Himalayan foothills to Kashmir (Kostrowicki, 1961). 

Clade31 [113(1), 142(1), 176(1), 177(1)]. There are no unique and unreversed synapomorphies uniting the 
seven constituent genera of clade 31. All four of the character states listed above are reversed in one or 
more genera, and three also occur in the outgroups. Characters 176 and 177 have been discussed in detail 
earlier; they are reversed in Loboplusia and Puriplusia respectively, and the former is also present in 
Brachionycha. A saccular sheet (142) is absent in Loboplusia, while convergent developments are to be 
found in Calophasia, Magusa and Lophoptera. Finally, the form of the vinculum (113) is interpreted as a 
synapomorphy for clade 31. However, the character is subsequently reversed in the included clade 35, and 
also occurs in Anagrapha. This, together with the uncertainty concerning the homology of states 1 and 2 of 
142 and the ambiguity provided by Loboplusia (and to a degree Anagrapha), suggests that clade 31 may not 
retain its current topology when further taxa and characters are analysed. 

Clade 32 [99(2), 137(1), 139(1), 191(1)]. Allagrapha and Macdunnoughia (s.l.) are interpreted as 
sister-taxa primarily on the basis of a produced tegumen apex (99). Also, the setae along the ventral edge of 
the valve are borne on marginal prominences (137), a condition otherwise only seen in Omorphina, where 
it is associated with a different valve shape. As noted earlier, clade 32 convergently shares a concealed 
clavus base (139) with Rachiplusia. A relationship between Allagrapha and Macdunnoughia (s.l.) was first 
suggested by Franclemont (1964). 

Allagrapha [3(0), 16(1), 18(1), 58(0), 127(0), 201(1)]. Allagrapha is here interpreted as a monotypic genus 
containing only A. aerea (Fig. 75) (McDunnough, 1944; Franclemont & Todd, 1983). Occasionally (e.g. 
Eichlin & Cunningham, 1978), a second species, aereoides (misspelt in McDunnough's (1944) figure 
captions, Eichlin & Cunningham (1978) and Franclemont & Todd (1983) as aeroides), has been included. 
Eichlin & Cunningham (1978) presented, as evidence for this inclusion, the absence in both species of a 
raduloid on the larval hypopharynx (present in Diachrysia balluca), modifications of the male genitalia and 
a strong superficial similarity. Ichinose (1973), however, synonymized Allagrapha (as Agrapha) with 
Diachrysia (as Plusia), noting a unique larval chaetotaxy (his type F) and the well-defined pocket in the 
female segment A8, discounting the modification to the sacculus in A. aerea (Fig. 368). 

However, while aereoides can remain in Diachrysia, I conclude that aerea is best retained for the present 
in the separate genus Allagrapha, on the basis of the characters discussed in previous sections. Neverthe- 
less, several Diachrysia species do complicate matters. D. leonina has a broad, cone-shaped vinculum 
(113); D. nadeja (Oberthiir) bears a small, broad, triangular spine on the sacculus (141); and the marginal 
setae on the valve of D. chrysitis, at least, are borne on small prominences (137). A more extensive analysis 
of Diachrysia may eventually show that not only is this genus more closely related to some Autoplusiina 
than to the Plusiina (as is concluded in the present analysis), but also that Allagrapha is merely a divergent 
species. However, it is unlikely that A. aerea and Diachrysia aereoides will prove to be sister-species. 

In addition to the characters already mentioned, Allagrapha also has several genitalic autapomorphies. 
The harpe is broad, flat and spoon-shaped; the clavus has a markedly swollen apex (Fig. 368); the LHS 
basal sclerotization on the vesica bears very strong, sharp spines; and the ostium bursae is protected by a 
large, spiculate lobe. 

Allagrapha aerea is an undistinguished brown plusiine, showing, as already noted, strong superficial 



156 IAN J. KITCHING 

similarity in wing shape and pattern to Diachrysia aereoides, D. leonina and D. bieti. It is widely distributed 
in southern Canada and the U.S.A., east of the Rocky Mountains (Eichlin & Cunningham, 1978). The 
larvae are generally polyphagous on low plants (Forbes, 1954) such as Urtica (Crumb, 1956), Aster and 
Glycine max (Papilionaceae; Eichlin & Cunningham, 1978). 

Clade 33: Macdunnoughia (s.l.) [145(1), 150(1), 192(1), 206(1), 213(3)]. The two constituent taxa of 
Macdunnoughia (s.l.), Macdunnoughia (s.str.) and Puriplusia, share two unique and unreversed synapo- 
morphies: a harpe in the form of an inwardly curving hook (150) and the presence of spinose pads lateral to 
the ostium bursae (192). Both characters are subsequently transformed, 150 in Macdunnoughia (s.str.) and 
192 in Puriplusia. In addition, and as previously noted, clade 33 convergently shares the apomorphic states 
of 145, 206 and 213 with Erythroplusiai 'Antoculeora, Rachiplusia and clade 35 respectively. 

Puriplusia [177(0), 192(2), 209(1)]. Puriplusia differs from Macdunnoughia (s.str.) primarily in the spinose 
pads lateral to the ostium bursae being elongated into recurved grooves (192). In addition, the apex of the 
corpus bursae has rotated through approximately 180 degrees and is thus directed anteriorly (209; see 
Character analysis). Puriplusia also lacks concave lateral edges to the female St7 (177), probably as a 
consequence of the reduced pleural membrane thickening (176). Autapomorphies not included in the data 
set are the saccular sheet that is modified into an incurved hook and a sacculus bearing a distinct, blunt 
spine (Fig. 369). The forewing pattern is also distinctive, although the high contrast two shades of grey seen 
in typical P. purissima is replaced by an almost uniform grey-brown, on which the cross-lines are barely 
distinguishable, in some forms of P. tetragona (Walker) (the synonymous P. semivitta (Moore); Dufay, 
1977). 

There are currently two species placed in Puriplusia, P. purissima (Fig. 77) and P. tetragona. P. semivitta 
may prove to be a third or merely a colour form of P. tetragona. However, the complex is in need of revision 
and there is at least one undescribed species in the BMNH collection. P. tetragona differs most markedly 
from P. purissima in the presence of a row of short, curved cornuti on the vesica, similar to those found in 
Macdunnoughia (s.str.) (see also Ronkay, 1982). Additionally, the recurved spiculate grooves lateral to 
the ostium bursae are less well developed and there is no black dash above the Y-mark, distal to the 
forewing discal cell. 

P. purissima occurs in China, Korea and Japan (Hampson, 1913). The larva was described by Ichinose 
(1960) but no record of the larval foodplant was made. Miyata (1983) gave Artemisia and Arctium 
(Asteraceae). P. tetragona is found further west and south in India. There are also specimens in the BMNH 
collection from Taiwan. The undescribed species referred to above is exceptional in being collected from 
west Sulawesi. 

Macdunnoughia (s.str.) [150(2), 158(2), 175(1)]. Macdunnoughia (s.str.) is characterized in this data set by 
the presence of an upward curving spine at the base of the harpe (150). It also bears a characteristic 
sharp-edged auricular cornutus at the base of the vesica (175) and, like Erythroplusia and the Argyrogram- 
matini, has the ventral sclerotization of the aedeagus restricted to a discrete narrow band (158). 

Four species were placed in Macdunnoughia (s.str.) (type species: M. confusa) by Kostrowicki (1961). 
Examination of the male genitalia of the holotype of M. camptosema Hampson showed that it lacks all the 
apomorphies listed above for the Autoplusiina and correctly belongs in Autographa (comb. n.). The male 
genitalia figured by Kostrowicki as A. camptosema (1961: fig. 54) are those of M. confusa(cf. his fig. 53). In 
addition, Ichinose (1973) has already pointed out that the female genitalia of M. crassisigna figured by 
Kostrowicki are actually also those of M. confusa. Thus, Kostrowicki, to whom the male of M. crassisigna 
and the female of A. camptosema were unknown, appears to have examined only one species of 
Macdunnoughia (s.str.) (the appropriately named M. confusal) but treated it as three separate species. 
The last species transferred to Macdunnoughia by Kostrowicki was M. schalisema (Hampson), although 
this time he did not examine any specimens. However, I have examined the genitalia of the female 
holotype of M. schalisema and conclude, from its lack of both lateral spiculate pads near the ostium bursae 
and a strongly ridged apex to the corpus bursae, that the species does not belong in Macdunnoughia. 
Instead, a comparison with the types of Autographa emmetra Dufay shows schalisema to be extremely 
similar both in forewing pattern and in female genitalia. I thus transfer M. schalisema to Autographa 
(comb, n.) and synonymize Autographa emmetra (syn. n.) with it. 

There are therefore only three species remaining in Macdunnoughia (s.str.): M. confusa (Fig. 76), M. 
crassisigna and M. monosigna Chou & Lu. The last of these closely resembles the second in male genitalia 
and may only be a colour form of M. crassisigna in which the basal half of the Y-mark has been lost. As 
noted above (150: Character analysis), M. crassisigna has only a rudimentary upcurving spine at the base of 
the harpe and is thus intermediate between M. confusa and Puriplusia. A more crucial feature, however, is 



SYSTEMATICS OF THE PLUSIINAE 157 

that in M. crassisigna, the apex of the corpus bursae is reflexed anteriorly and elongate, as in Puriplusia 
purissima. If this condition is interpreted as apomorphic, and a comparison with the other members of the 
Autoplusiina suggests that it is, then a sister-group relationship between M. crassisigna and Puriplusia is 
indicated, rendering Macdunnoughia (s.str.) paraphyletic. 

M. confusa is a trans-Palaearctic species ranging from France, across central Europe and Asia, to 
northern China, Japan and even the southern tip of Kamtchatka (Kostrowicki, 1961). However, until 
about 1918, the species was only known from eastern Europe. After this date, M. confusa began a rapid 
westward expansion of its range and is now probably resident in Italy, Spain and France. The species was 
first recorded from Great Britain in 1951 (Dewick, 1951) but never became resident. It appeared to have 
reached its limits in western Europe by 1955 (Bretherton et al. , 1983). 

M. crassisigna occurs in China, the Korean peninsula and Japan, extending westward through northern 
Burma to India and Pakistan (Kostrowicki, 1961), while M. monosigna has only been recorded from 
somewhere in China. 

The larvae of both M. confusa and M. crassisigna have been described by Ichinose (19586 and 19626 
respectively). Both species appear to be somewhat polyphagous, but with a slight bias towards members of 
the Asteraceae (Beck, 1960; Kostrowicki, 1961; Ichinose, 19626; Miyata, 1983). Oord (1985) stated that 
M. confusa larvae fed on Artemisia before hibernation and Lactuca and Matricaria after reawakening in 
spring. 

Clade 34 [12(1), 59(0), 148(0)] . The three character states that unite the four constituent genera of clade 34 
are very weak and it is unlikely that future studies will corroborate its monophyly. The main problem is 
Loboplusia (q.v.) with its highly simplified male valves and numerous missing data. This genus is probably 
not closely related to clade 35 (Sclerogenia, Autoplusia and Antoculeora). 

Loboplusia [6(1), 141(1), 142(0), 144(0), 175(1), 176(0), 209(1)]. Loboplusia is perhaps most notable in the 
present data set for the autoplusiine structures it does not possess. The genus lacks a broad saccular spine 
(141), a saccular sheet (142), has a reduced clavus( 144) (Fig. 371) and, while the lateral edges of St7 in the 
female are probably concave (177), the pleural membrane is unmodified (176). The genus possesses an 
auricular cornutus (175), but as the vesica of the slide preparation was not everted, its position on the vesica 
is only assumed to be basal . Also , like Puriplusia , the apex of the corpus bursae is rotated to face anteriorly. 

Nevertheless, the male Loboplusia has the most remarkable array of autapomorphic scent organs, their 
diversity exceeding even Chrysodeixis. In addition to the usual sternal hair pencils, the hind tibia is 
considerably swollen and bears a dense dorsal fringe of long, golden hairs. The hind tarsi are similarly 
adorned but are of normal dimensions. The anal angle of the hindwing is extended into a long lobe 
(superficially similar to the anal scent fold of swallowtail butterflies), which is folded and contains 
numerous long hairs. Finally, the outer edge of the hindwing upperside and the basal costal edge of the 
hindwing underside also bear modified scales. All in all, the male Loboplusia is a very distinctive and 
unmistakable plusiine. In contrast, the female lacks all these features and has a similar appearance to an 
Antoculeora, although without its golden scaling. 

I know of only five specimens of the sole species contained in Loboplusia, L. vanderweelei (Fig. 74); 
three males and two females. The holotype and paratype males were collected at 1700 m in west Java 
(Roepke, 1941). Subsequently, Dufay (1973) described two females as a separate species, Autographa 
pokhara. These were taken at 1500 m and 2000 m in Nepal and Burma respectively. The sexes were finally 
associated by Dierl (1975), who reported a third male, which he had taken in Nepal at 2360 m, near 
the locality of the holotype of A. pokhara. Little is known of the biology of the species, although Dierl 
(1975) stated that it is only to be found in very wet mountain forests receiving over 6000 mm of rain per 
year. 

Clade 35 [113(0), 142(2), 198(1), 212(1), 213(3)]. The three genera of clade 35, Sclerogenia, Antoculeora 
and Autoplusia are united by three good synapomorphies: the fusion of the saccular sheet with an overfold 
of the costa (142); increased sclerotization posterior to the ostium bursae (198); and differential 
sclerotization of the corpus bursae granulations (212). They also display strong sclerotized ridges on the 
apex of the corpus bursae (213), a feature also seen in Macdunnoughia (s.l.). 

Sclerogenia [95(0), 108(1), 155(1)]. Sclerogenia is another monotypic genus. The sole species, S. Jessica 
(Fig. 78), has several unique features of the genitalia not included in the current data set. The juxta has a 
very pronounced basal bulge, while the fused saccular sheet/costal overfold forms an elongate, ventrally 
curved process that just exceeds the bulged ventral edge of the valve (Fig. 361). The corpus bursae of the 
female is distinctive (Fig. 455) in that the apex is reflexed and fused along the posterior section of the fundus 



158 IAN J. KITCHING 

and is entirely open to it internally, thus giving the appearance of two strigate ventral sacs. The remainder 
of the fundus is characteristically lemon-shaped. 

Sclerogenia was initially erected as a subgenus of Autographa by Ichinose (1973) to receive the species 
Jessica, previously included in Macdunnoughia (Ichinose, 19626). Ichinose (1973) still considered S. Jessica 
to be closely related to Macdunnoughia (then also a subgenus of Autographa), in view of the similarities of 
the female genitalia. Sclerogenia was subsequently elevated to full generic status by Sugi (1982). 

S. Jessica is another species found in the eastern Palaearctic and southern Himalayan foothills. Hampson 
(1913) recorded the species from Japan, Assam and the Punjab, while it also occurs in China (Chou & Lu, 
1978). There is also a single specimen in the BMNH collection from northern Luzon in the Philippines. The 
larva of 5. Jessica was described by Ichinose (1960a). Its larval foodplants appear to be restricted to 
members of the Asteraceae: Sonchus, Lactuca (Ichinose, 1973), Ixeris and Arctium (Miyata, 1983). 

Clade 36 [158(0), 198(2), 199(1), 200(1)]. Antoculeora is interpreted as the sister-group of Autoplusia on 
the basis of three synapomorphies: further sclerotization of the ventral area posterior to the ostium bursae 
(198); a ventral origin of the anterior apophyses (199); and a distinctive form of antrum (200). In addition, 
the aedeagus sclerotization of both genera is not restricted to a ventral band, instead the organ is generally 
sclerotized, as in the presumed plesiomorphic state. 

Antoculeora [18(1), 104(1), 116(0), 117(0), 118(2), 145(1), 189(0), 191(1)]. The outstanding features of the 
genus Antoculeora are the highly modified and very distinctive genitalia. In the male, the whole of the 
genital capsule appears to have moved posteriorly and to have rotated somewhat dorsally. This has 
resulted in the pleurites losing contact with the dorsal edge of the tegumen arms, only retaining apical 
contacts (118), and straightening (116) and narrowing (117) in the process. The tegumen arms are 
extremely swollen anteriorly and, in dorsal view, diverge at 180 degrees (104), leaving a broad, U-shaped 
anterior emargination . The valves have become irregularly and strongly digitate , taking the form of a stag's 
antlers (Figs 376, 377). The tines are irregularly curved and asymmetrical, the left valve having three, the 
right four, so that they partially interlock when the valves are closed. The saccular sheet has become a long, 
apically serrate lance. The modifications in the female are less extreme but are doubtless partly correlated 
with those of the male. St7 is medially and posteriorly deeply emarginate, while the antrum has become a 
heavily sclerotized, open, irregularly and asymmetrically lobed chamber (Figs 436, 437, 456). Together, 
they form a unique genital complex within the Plusiinae. 

There are three, perhaps four, very closely related species in Antoculeora: A. ornatissima (Fig. 79), A. 
locuples, A. lushanensis Chou & Lu and A. minor Chou & Lu, differing only in minor details of wing 
pattern and valve shape. They may all eventually be referable to a single species. Initially, A. ornatissima 
was placed by Kostrowicki (1961) in Argyrogramma. This, however, was based upon a misidentification 
(Ichinose, 1963a; Dufay, 1970; Ichinose, 1973). Ichinose (19626) included ornatissima tentatively in 
Macdunnoughia, then erected Antoculeora to accommodate it (Ichinose, 1973). Antoculeora was first 
treated as a subgenus of Erythroplusia but a relationship with Macdunnoughia was also suggested 
(Ichinose, 1973). This was formalized by Dufay (1977), who transferred ornatissima and locuples (as a 
synonym of the former) to Macdunnoughia. Antoculeora was resurrected as a full genus by Chou & Lu 
(1979a) and so treated by Sugi (1982). 

A. ornatissima has a similar distribution to Sclerogenia: Japan, Korea, China, the far eastern U.S.S.R. 
(Primorskiy Kray), Sikkim, Punjab, Nepal and Bengal (Dufay, 1977). A. minor and A. lushanensis are so 
far only reported from China, while A. locuples is recorded in the BMNH collection from Japan and the 
U.S.S.R (Primorskiy Kray). Nothing is known about the biology of this genus. Ichinose (1973) stated that 
the larvae had been reared on Sonchus and Lactuca but that the natural foodplant was unknown, while 
Miyata (1983) gave Petasites. If these prove correct, then Antoculeora displays the same oligophagy, 
restricted to the Asteraceae, as its close relative Sclerogenia. 

Autoplusia [12(0), 14(2), 59(1), 64(1), 83(3), 131(0), 140(0), 216(1)]. Autoplusia is an exclusively New 
World genus that is in need of revision. Four species are currently recognized: A. egena, A. olivacea, A. 
egenoides Franclemont & Todd and A. illustrata (Guenee). To those can be added 'Syngrapha' gammoides 
and I will limit my discussion to these five species. However, I understand from Mr F. Navarro that there 
are several more species, currently languishing in either Syngrapha or 'Plusia' (s.l.) that are assignable to 
Autoplusia. 

A. egenoides was originally described by Hampson (1913) as the 'much darker' 'ab. 1' of A. egena. It was 
named 'ab. egenoides' by Strand (1917), a name employed subsequently by Draudt (1940). For some 
mysterious reason, it was elevated to full species by Franclemont & Todd (1983) in the latest North 
American checklist. Even the very inclusion of A. egenoides in that list is odd for, as far as I am aware, only 



SYSTEMATICS OF THE PLUSIINAE 159 

the holotype exists and that was collected in Mexico. Nevertheless, it would appear that egenoides has been 
wrongly associated from the outset. I have examined the holotype and conclude that A. egenoides is 
probably synonymous with A. olivacea. However, comparison of the genitalia is required to confirm this. 

Autoplusia has a number of features in common with Rachiplusia, as noted above (characters 14, 64 and 
216). It is also the only genus in the Plusiini to lack a saccular flange. In addition, as stated above, while the 
type species, A. egena, lacks a broad, triangular saccular spine (Fig. 372) (141), this structure is present in 
A. olivacea (Fig. 373) and 'Syngrapha' gammoides (Fig. 374). The autoplusias also possess a number of 
genitalic synapomorphies. The saccular sheet/costal overfold fusion is extended into a curved hook, the 
apex of the valve forming an oval lobe arising from the external surface of this structure. The sacculus is 
extremely inflated. The everted vesica curves at about half its length through 90-200 degrees, the angle 
varying between individuals within a species. On the outer surface of the angle, there is a large, distally 
directed, thorn-like cornutus, while the apical half of the vesica is covered with numerous small, 
basally-directed cornuti, the arrangement of which is species specific. The distal part of the ductus bursae is 
heavily sclerotized, dorso-ventrally flattened and strigate, while the apex of the corpus bursae is elongated 
and apically recurved (Fig. 454). These features, with modifications, are present in A. egena, A. olivacea 
and 'Syngrapha gammoides. 

A. illustrata, however, lacks all the valve apomorphies listed above but may have the thorn-like cornutus 
(Eichlin & Cunningham, 1978: fig. 59). The female genitalia also appear to share few features in common 
with the other members of the genus (Eichlin & Cunningham, 1978: fig. 95) , while close examination of the 
forewing pattern reveals only a superficial similarity. A. illustrata was only tentatively included in the genus 
by Kimball ( 1965) and although Eichlin & Cunningham ( 1978) stated that the male genitalia supported this 
decision, I disagree. A detailed study of Autoplusia will probably show that A. illustrata (and the similar 
'Syngrapha' egenella (Herrich-Schaffer)) ought to be removed from this genus. At present, I cannot 
suggest a better placement. 

'Syngrapha' gammoides (Figs 82, 83) is a Chilean endemic. Its life history and immature stages have been 
described in detail by Angulo (1973) and Angulo & Weigert (1975). It is recorded as feeding on several 
crop plants in Chile and is an economically important pest in that country. A. egena is found from the 
southern U.S.A. to Brazil and is also a pest species and high polyphagous (Eichlin & Cunningham, 1978). 
It is commonly known in the U.S.A. as the Bean Leaf Skeletonizer and there have been occasional serious 
outbreaks of this species (Kimball, 1965). The larvae have been described by Crumb (1956) and Eichlin & 
Cunningham (1978). 

The last species, A. olivacea (Fig. 81), was stated by Eichlin & Cunningham (1978) to be found only in 
southern coastal California. However, in the course of this study, I found that there has been considerable 
confusion between this species and A. egena (Fig. 80). The most obvious wing pattern difference concerns 
the form of the terminal line between the forewing veins R 5 and M 3 . In A. egena, it forms a distinct, single 
pink crescent (Fig. 80), while in A. olivacea, the terminal line in this section of the wing is less distinct and is 
broken into three small crescents, one between each pair of veins (Fig. 81). Additionally, in A. egena, the 
forewing postmedial line is a distinct, thin, brown line, which is flanked on its outer edge by a broader, 
pinkish brown band. It passes diagonally across the wing towards the apex but recurves basally just before 
it meets the costal margin (Fig. 80). In A. olivacea, only the pinkish brown band is present. This becomes 
very indistinct apically and does not recurve to meet the costa (Fig. 81). The Y-marks in both species are 
very variable and of no use in identification. Those illustrated in Figs 80, 81 represent extreme forms that 
occur in both species. These forewing pattern elements coincide with the much more distinctive genitalic 
differences (see Eichlin & Cunningham, 1978). From this discovery, I draw three conclusions. First, the 
specimens illustrated by Eichlin & Cunningham (1978: figs 168, 170) as A. olivacea and A. egena are both 
A. olivacea. Second, A. olivacea is much more widespread than was thought. Specimens in the BMNH 
collection have been taken in California, Mexico, Guatemala, Costa Rica and Colombia. Third, overall 
forewing ground colour is of little use in distinguishing the two species. While A. olivacea, when fresh, is 
distinctly darker and greener than A. egena, both fade to a similar pale brown. Incidentally, the holotype of 
A. egenoides has the forewing pattern of A olivacea, hence my suggestion above that the two species may 
prove to be synonymous. The larva of A olivacea has been described by Eichlin & Cunningham (1978) 
and, like A. egena, is polyphagous. 

Clade 37: Plusiina [83(1), 146(1)]. The constituent genera of the subtribe Plusiina are united solely by the 
apomorphic state of character 146, a blade-like valve. As already noted above (Characters), this character 
state is difficult to define but is readily recognized. The condition may have been misinterpreted in 
Anagrapha (q.v.). 

Diachrysia [3(0), 9(1), 16(1), 58(0), 65(1), 165(1), 191(1)]. Ten species are currently placed in the Holarctic 



160 IAN J. KITCHING 

genus Diachrysia. Two, D. aereoides and D. balluca, are Nearctic, the remainder Palaearctic, with a 
distinct concentration in Japan and neighbouring countries. As stated earlier, Diachrysia may not be 
monophyletic, particularly with regard to Allagrapha, and thus the sole autapomorphy included in the data 
set (65: 1 ; the presence of a small number of large SD and D tarsal spines) may not be found much beyond 
D. chryson and D. coreae Ogata). 

There appear to be three species-groups within Diachrysia. The chryson species-group comprises five 
species: D. aereoides, D. leonina, D. bieti, D. chryson (Fig. 84) and D. coreae. D. aereoides differs in being 
a small, paler brown moth than the others. It is widespread in the northern U.S.A. and southern Canada, 
extending south in the west to Colorado and in the east to Pennsylvania (Eichlin & Cunningham, 1978). 
The larvae are recorded as feeding upon Mentha (Lamiaceae), Spiraea (Rosaceae) (Crumb, 1956) and 
Aster (Forbes, 1954), and have been described by Crumb (1956) and Eichlin & Cunningham (1978). The 
other four species are among the largest plusiines and have somewhat falcate forewings. D. bieti and D. 
leonina are closely related and have brown forewings without any extensive metallic markings. The larva of 
D. leonina has been illustrated by Kogi (1986). It bears enlarged conical dorsal pinacula, very similar to 
those seen in the larva of Anadevidia. D. leonina is recorded as being polyphagous on such genera as 
Urtica, Aster and Cirsium. D. coreae is similar to D. bieti and D. leonina but bears a diffuse, subapical, 
quadrate, golden-green metallic patch on the forewing. It has long been regarded as synonymous with D. 
chryson (e.g. Kostrowicki, 1961), in which the patch is much more discrete and intense. Nothing is 
recorded of the biology of D. coreae but that of D. chryson is well known (e.g. Haggett, 1958; Bretherton er 
al. , 1983). The chaetotaxy has been described by Ichinose (1962£>). D. chryson appears to be facultatively 
monophagous on Eupatorium (Asteraceae) (e.g. Bretherton et al., 1983) but other foodplants have 
been recorded: Salvia, Mentha (Lamiaceae) (Kostrowicki, 1961) and Arctium (Asteraceae) (Ichinose, 
19626). 

The zosimi species-group comprises only D. zosimi, and possibly D. balluca, although this latter species 
also shows some resemblances to the chry son-group. D. zosimi has a unique colour pattern, in which the 
forewings dorsal and distal of the discal cell are entirely metallic golden-green, with a 'crushed velvet' 
effect. The biology of this trans-Palaearctic species (Kostrowicki, 1961) has been described in detail by 
Kasy (1953). The larvae are polyphagous but mostly on species of Rosaceae (Kasy, 1953; Kogi, 1986). D. 
balluca has a similar forewing pattern to D. zosimi but has the large size and falcate forewing of the 
chry son-group. It is restricted to the north-eastern U.S.A. and south-eastern Canada. The larval 
chaetotaxy has been described by Eichlin & Cunningham (1978) and the foodplants have been given as 
Humulus (Cannabinaceae) (Saunders, 1873) and Populus (Eichlin & Cunningham, 1978). 

The third species-group, exemplified by D. chrysitis (Fig. 85), comprises three species in which the 
forewing pattern consists of an H-shaped metallic patch, which may occasionally be represented by two 
separate, transverse bands instead. D. stenochrysis (Warren) has a more diffuse pattern than the other two, 
D. chrysitis and D. nadeja, and is restricted to the far eastern Palaearctic (Kostrowicki, 1961). The genitalia 
of D. stenochrysis can be difficult to distinguish from those of D. chrysitis, especially the males (Ronkay, 
1982) . D. nadeja was also thought to occur only east of the Ob' River basin , with isolated records as far west 
as Bessarabia (Kostrowicki, 1961). However, like Macdunnoughia confusa, D. nadeja appears to be 
undergoing a rapid westward expansion of range, having been recently discovered in Austria (Aistleitner 
& Rezbanyai, 1982), Switzerland (Rezbanyai, 1980) and France (Dufay, 1986). If this progression 
continues, then it may only be a matter of time before D. nadeja is captured in Great Britain and 
Scandinavia. The life history has been described by Rezbanyai (1983), with tftat of D. chrysitis for 
comparison. The natural foodplants of D. nadeja are unknown but Urtica and Plantago (Plantaginaceae) 
were accepted in captivity (Rezbanyai, 1983). 

D. chrysitis is a very variable species, as evidenced by no fewer than 14 varietal and aberrational names. 
The species is widespread across the Palaearctic from Ireland to Japan. The immature stages have been 
described in detail by Rezbanyai (1983) and numerous other accounts exist (e.g. Bretherton et al. , 1983). 
The main larval foodplant is Urtica but many other species can be utilized. 

There is also an occasionally recurrent taxonomic problem concerning D. chrysitis and the supposed 
species D. tutti (Kostrowicki). D. tutti was described as a separate species from D. chrysitis (Kostrowicki, 
1961) on the basis of smaller size and minor differences in forewing pattern and in the genitalia (Bretherton 
et al. , 1983). Lempke (1965) examined a long series of Dutch specimens and agreed that tnere were two 
species, although there was also a great deal of intraspecific variation. However, Lempke (1966a) 
subsequently re-examined some of his material and found that, with the aid of two pins, he could transform 
the narrow ostium bursae [= antrum] of D. chrysitis into the broader ostium bursae of D. tutti. He thus 
concluded that he had been wrong and that there was, after all, only one species, D. chrysitis, which was 
subject to geographic variation (not unexpected given the extensive range of the species) . Bretherton et al. 
(1983) regarded tutti merely as an aberration of D. chrysitis. However, Rezbanyai (1985) has recently 



SYSTEMATICS OF THE PLUSIINAE 161 

suggested that tutti may represent an eastern subspecies of D. chrysitis that is only partially reproductively 
isolated from the nominate subspecies by incomplete pheromonal differences. The two taxa, which have 
subsequently become partially sympatric, are thus currently in the process of hybridizing, further clouding 
the distinction between them. The problem is obviously not yet fully resolved. 

Clade 38 [105(1)]. This clade is supported by only a very homoplasious character state and is unlikely to be 
corroborated by the results of future analyses. 

Anagraphs [64(2), 113(1), 127(2), 143(2), 144(1), 189(1)]. Anagrapha is yet another of those genera that 
shares putative synapomorphies with two widely separated groups of genera on the cladogram. The genus 
is usually associated with Syngrapha (s.l.), due to the spined hind tibiae (64) and certain larval characters 
(see below). They also share a juxta with a subapical process (127) and a T8 that is irregularly cut back 
ventral to the base of the anterior apophysis (189). However, this latter character state also occurs in the 
Autoplusiina, while a broad, flat, cone-like vinculum (113) further supports a relationship between some 
members of this tribe and Anagrapha. Lastly, there is a remarkable similarity between Anagrapha and 
Macdunnoughia (s.str.) (especially M. confusa) in forewing pattern (cf. Figs 86, 76 respectively). 

However, Anagrapha shows none of the genitalic features characteristic of Macdunnoughia and its 
relatives, although it has a number of autapomorphies. The harpe is bulbous basally and apically, and is 
bent outwards at 90 degrees at about half its length. The valve is deeply excavated apically and the costa has 
a fringe of dense, persistent, yellow hair. The costa terminates in a single distinct spine while the distal part 
of the sacculus bears broad, curved setae, somewhat like those found in Ctenoplusia and its relatives. In the 
female, the apex of the corpus bursae is recurved, as in many autoplusiines, but, while heavily sclero;.zed, 
lacks the characteristic strong internal ridges of that subtribe (Fig. 459). 

Anagrapha was erected by McDunnough (1944) to accommodate the species A. falcifera (Fig. 86). With 
its markedly different genitalia, the species was considered to be a highly divergent branch of Syngrapha. 
Eichlin & Cunningham (1978) agreed with this interpretation and synonymized Anagrapha with Syn- 
grapha, tentatively deriving A. falcifera from the Syngrapha diasema lineage. One of the main reasons for 
their placing of Anagrapha was the presence of seta SV2 on segment Al of the larva. The feature was used 
to unite Syngrapha, Anagrapha, Plusia and a branch of Autographa. However, while this seta also occurs in 
Sclerogenia and Macdunnoughia (s.str.) (Ichinose, 19626), tantalizingly, it is absent in all other members 
of the Autoplusiina known so far, and most notably in Puriplusia and Allagrapha. Thus, the relationships 
of Anagrapha are as problematical using larval characters as with using adult characters and they are still 
far from clear. Anagrapha was subsequently reinstated to full generic status by Franclemont & Todd 
(1983). 

A. falcifera is an abundant species throughout the U.S.A. and southern Canada (Eichlin & Cunning- 
ham, 1978). No records are known for the countries south of the U.S.A. Commonly known as the Celery 
Looper, the larvae sometimes cause economic losses to crops (Forbes, 1954). Overall, however, A. 
falcifera appears to be of only minor agricultural importance. 

Clade 39 [12(1), 108(1), 109(1)]. The most characteristic feature of clade 39 is the strongly S-shaped 
vinculum, when seen in lateral view. However, this state is also weakly expressed in clade 21 of the 
Euchalciina and further evidence is required to support the monophyly of clade 39. 

Autographa gamma [18(0)]. Autographa gamma (Fig. 87) was chosen to represent that branch of the genus 
characterized by the lack of seta SV2 on segment Al of the larva (Eichlin & Cunningham, 1978). It is thus 
part of an overtly paraphyletic group of uncertain limits (cf. Trichoplusia). Eichlin & Cunningham 
included ten species of Autographa in this group, divided into three lineages, depending upon the number 
of processes on the ribs of the larval mandible. 

The first lineage, A. precationis (Guenee) and A. bimaculata Stephens, was interpreted as the most 
primitive because the processes on both ribs 2 and 3 had been retained. McDunnough (1944) noted that A. 
precationis was unusual and displayed a number of features in common with the Argyrogrammatini, 
notably a bulbous coecum (157) and apical, basally-directed, setose scales on the valve (136). 

The next most derived lineage had lost the process on the third mandibular rib and also comprised two 
species, A. sansoni and A. rubida Ottolengui. 

Those species that had lost both mandibular processes were considered to be the most derived and 
included six Nearctic species: A. biloba, A. californica, A. pseudogamma (Grote), A. v-alba Ottolengui, 
A. pasiphaeia (Grote) and A. flagellum (Walker). However, Neil (1984) has shown that the larva of A. 
flagellum, which was unknown to Eichlin & Cunningham (1978), does possess a process on mandibular rib 
2, and thus may be more closely related to A. sansoni and A. rubida. 



162 IAN J. KITCHING 

Of the Palaearctic species, five were examined from specimens in the BMNH collection, of which A 
gamma was found to lack SV2 on Al (so too did A. bractea, see below). 

Of the species for which life histories have been elucidated, most have been shown to be polyphagous on 
herbaceous plants. A few (A. bimaculata, A. pasiphaeia, A. flagellum) have been recorded from only one 
or two genera of larval foodplants (Eichlin & Cunningham, 1978), but this may reflect lack of information 
rather than true oligophagy. 

Two species are serious agricultural pests; A. gamma in the Palaearctic (Carter, 1984 and references 
therein) and A calif ornica in the western Nearctic (Hyslop, 1912). A. biloba is an occasional pest of garden 
crops (Forbes, 1954). 

Clade 40 [6(0), 215(1)]. The members of clade 40 are united by the synapomorphic possession of a lateral 
patch of thickened membrane on the corpus bursae (215). This feature is, however, lost in clade 44 
(Syngrapha s.l.) and this must weaken the expectation that clade 40 will be corroborated by future studies. 

Clade 41 [215(2)]. Autographa bractea and Cornutiplusia are united as sister-taxa by the further develop- 
ment of the lateral thickened membrane on the corpus bursae into at least a shallow pouch (215). This 
structure also occurs in the presumed close relatives of A. bractea (see below), A. aemula and A. excelsa 
(Kretschmar) (Kostrowicki, 1961). 

Autographa bractea [6(0), 12(0)]. A. bractea (Fig. 88) was originally chosen to represent the second 
Autographa lineage comprising species that possess seta SV2 on larval segment Al. The choice was made 
by picking the British equivalent of an American species (A. metallica (Grote)) included in the second 
lineage by Eichlin & Cunningham (1978). However, they had not examined the larva of A. metallica and 
when, subsequent to my decision, I examined the larva of A. bractea, I found that it actually lacked seta 
SV2 on Al. A. bractea (and presumably those species, including A metallica, that share the distinctive 
forewing pattern; see below) is thus referable to the presumed plesiomorphic (paraphyletic) lineage of 
Autographa exemplified by A. gamma. I will therefore discuss two further subgroups of Autographa, A. 
bractea and its relatives, and those species known to possess seta SV2 on Al . 

A. bractea could be considered to be part of a species-group in which the Y-mark has become modified 
into an enlarged, single, silver or gold spangle. There are five species with such a pattern: A. bractea, A. 
metallica, A. aemula, A. excelsa and A kostjuki Kljutshko. The group is Holarctic in distribution, although 
A metallica is the only American representative. The life histories of three are known: A aemula 
(Hirschke, 1898), A excelsa (Laasonen & Skvortsov, 1984) and A. bractea (e.g. Beck, 1960; Brethertone? 
al. , 1983). All three species are polyphagous but with a preference for the Asteraceae. 

There are six species currently placed in Autographa that are known to have seta SV2 on segment Al: A. 
corusca, A. mappa, A. ampla (Eichlin & Cunningham, 1978), A. nigrisigna (Walker) (Ichinose, 1958a, b), 
A. ]ota and A pulchrina (Kitching, unpubl. data). The first three appear to feed as larvae primarily on 
woody plants (Eichlin & Cunningham, 1978), although only A ampla may be adequately recorded. While 
the other three are polyphagous on herbaceous plants (Beck, 1960; South, 1961 ; Miyata, 1983), A jota will 
also feed on unspecified deciduous trees and shrubs (Bretherton et al, 1983). A nigrisigna is an 
agricultural pest in the eastern Palaearctic, where it is known as the Asiatic Common Looper or the 
Cabbage Semilooper. There have been numerous studies of its bionomics (Ichinose, 1959; 1960ft; Ichinose 
& Asawa, 1964; Ichinose & Shibuya, 1959), while the chaetotaxy of the larva must be the most 
over-described of any plusiine, this having been presented in detail in at least three separate papers 
(Mukerji & Singh, 1951; Ichinose, 1960a; Betala & Kushwaha, 1972). 

As was noted for Thysanoplusia, delimiting a group based on larval characters, when the larvae of only a 
few species are known, results in many species possibly being wrongly excluded. There are several 
autographas that fall into this category, and the elucidation of their life histories is an important 
prerequisite to a comprehensive revision of the genus. 

Cornutiplusia [64(1), 215(3)]. In Cornutiplusia, the lateral pouch of the corpus bursae mentioned above 
(clade 41 ) is produced into a large pouch or into an appendix. The degree of development of this structure is 
geographically variable and is the subject of another study (Kitching, in prep.). Cornutiplusia also has 
spined hind tibiae and was thus associated with the genus Syngrapha (Hampson, 1913). Kostrowicki 
(1961), however, thought the general appearance and genitalia suggested a closer link with Autographa. 
Cornutiplusia stat. rev. is here treated as a good genus, certainly distinct from Syngrapha, perhaps less so 
from Autographa (s.l.). 

Cornutiplusia comprises a single species, C. circumflexa (Fig. 89), found throughout the more arid areas 
of southern Europe, North Africa and western Asia, as far as Sri Lanka and Lake Balkhash (U.S.S.R.). 



SYSTEMATICS OF THE PLUSIINAE 163 

The species extends down the African rift valley to Kenya and Uganda, and there are isolated populations 
on Grand Comoro and in South Africa/Namibia and the Gambia. The life history and larva have only been 
very briefly described (Kirby, 1907). The larval foodplants have generally been stated to be members of the 
Solanaceae (Kirby, 1907; Kostrowicki, 1961; Pinhey, 1975, who also gave Eschscholzia [Papaveraceae]). 

Clade 42 [166(1)]. The three taxa that constitute clade 42 (Plusia, Lophoplusia and Syngrapha (s.l.)) are 
united by their apparent shared possession of a grooved subapical cornutus. However, as discussed above 
(648-step cladograms), the presence of such a structure has yet to be confirmed in Syngrapha (s.l.). The 
species examined in this study all lacked a true subapical cornutus and were coded 'not comparable'. The 
results of future studies may therefore show that character 166 is actually a synapomorphy for Plusia and 
Lophoplusia. 

Plusia [12(0), 58(0), 83(3), 105(0)]. Plusia is a small genus containing at least five good species and a 
number of other dubiously distinct taxa. All except one (P. venusta Walker) have a diagnostic forewing 
pattern consisting of a yellow to deep orange ground colour, with the Y-mark modified into two large, 
silver spangles and a white dash present in the cell M\ , between the discal cell and the subterminal line (Fig. 
90). In P. contexta Grote, these markings have become somewhat elongate, a transformation series that 
culminates in P. venusta (Fig. 91). In this species, the Y-mark has become a thin, white streak, while the 
ground colour of the forewing has become pale brown with cream margins. It thus superficially resembles 
other noctuid genera such as Mythimna Ochsenheimer and Simyra Ochsenheimer. All species of Plusia 
also have reduced dorsal abdominal tufting. 

The species are all very similar in their genitalia. The males of P. venusta, P. putnami Grote and P. 
nichollae (Hampson) cannot be reliably distinguished, while the females of the first two are similarly 
virtually identical. P. festucae differs from P. putnami on minor points of genitalic structure, such as having 
a larger clavus, a longer subapical cornutus that is pointed at both ends and a shorter ductus bursae 
(Lempke, 19666; 1969; Dufay, 1969; Bretherton et al. , 1983). The males of P. contexta are unique in that 
the valve costa terminates in a narrow, blunt projection (Eichlin & Cunningham, 1978). P. festucae and P. 
nichollae differ from the other three species in having a somewhat round sclerotization on the corpus 
bursae rather than one that is elongate. 

The genus Plusia is Holarctic. P. festucae (Fig. 90) occurs across the Palaearctic region from the British 
Isles to Japan. It is highly variable in size and wing pattern and this has led to the naming of numerous 
aberrations, varieties and parochial subspecies of doubtful reality. P. putnami is even more widespread and 
has a similar number of subspecific and infrasubspecific names. Four subspecies are currently recognized: 
P. p. putnami occurs west across the northern U.S.A. and southern Canada to the Rocky Mountains but is 
absent from the central states (cf. Pseudeva purpurigera) (Eichlin & Cunningham, 1978); P. p. gracilis 
(Lempke) is restricted to western Europe (Dufay, 1969; Bretherton etal. , 1983); P. p. barbara (Warren) is 
known only from a unique female holotype from Morocco; and P. p. festata Graeser occurs in the eastern 
Palaearctic, from the Ob' River basin to Japan. The remaining three species are all North American. P. 
venusta and P. contexta have similar ranges to P. p. putnami, although neither extends as far west, while P. 
nichollae is restricted to the west coast, from northern California to southern Alaska. 

Larval foodplants are known for all species except P. nichollae. The genus Plusia feeds primarily on 
grasses, sedges and other marsh and waterside monocotyledons (Beck, 1960; Dufay, 1969; Eichlin & 
Cunningham, 1978; Bretherton et al., 1983; Miyata, 1983). The last group includes Sparganium (Sparga- 
niaceae), Typha (Typhaceae), Iris (Iridaceae), Alisma (Alismataceae) and Juncus (Juncaceae). P. festucae 
has also been reared on Crocosmia (Iridaceae), Salix, Brassica and Linum but these are species accepted in 
captivity, rather than the natural foodplants. The larva shows several features unique in the Plusiinae that 
are probably linked to its monocotyledonous diet. It is elongate, slender and cylindrical (Bretherton etal. , 
1983) and the head is about as wide as the rest of the body. The larva does not, therefore, taper anteriorly as 
is typical of other plusiines. The crotchets are uniordinal, while the mandibles have a deep pocket on the 
inner ventral edge that is apparently a modification for feeding on grasses (Eichlin & Cunningham, 1978). 
Pupation occurs in an elongate, strong, white cocoon, spun vertically on the underside of the leaves of its 
foodplant and is placed near the apex so as to cause the leaf tip to droop and conceal it (South, 1961). 

Clade 43 [59(0), 144(1), 166(2)]. There are no convincing synapomorphies at present to unite Lophoplusia 
and Syngrapha (s.l.) as sister-groups. Characters 59 and 144 have been shown to be highly unreliable. The 
possibility that a grooved or winged subapical cornutus (166) actually represents a synapomorphy for 
Lophoplusia and Plusia, to the exclusion of Syngrapha (s.l.), has been discussed above. 

Lophoplusia [18(0), 46(2), 47(1), 65(3), 107(1), 144(2)]. The genus Lophoplusia is endemic to the 



164 IAN J. KITCHING 

Hawaiian Islands. There are currently four described species: L. pterylota (Fig. 92), L. psectrocera, L. 
violacea (Swezey) and L. giffardi (Swezey), with a possible fifth undescribed species represented in the 
BMNH collection. The most readily recognizable feature of all species is the bipectinate antennae of both 
sexes (46, 47) and this was the character state used by Zimmerman (1958) to define Lophoplusia when first 
described. In addition, L. pterylota has numerous long, fine SD and D tarsal setae (65) and a single, dorsal 
bar on the anal tube (107). L. giffardi shares the distinctively winged subapical cornutus (Zimmerman, 
1958: fig. 303) but its presence in the other species requires confirmation. 

Little is known about the biology of Lophoplusia. L. violacea and L. pterylota have recently been reared 
on Stenogyne (Lamiaceae) and Clermontia (Lobeliaceae) respectively (K. Sattler, pers. comm.). 

Clade44: Syngrapha (s.l.) [64(1), 97(1), 127(2), 149(1), 162(0), 189(1), 191(1), 201(1), 215(0)]. Syngrapha 
(s.l.) (i.e. including Palaeographa and Caloplusia, but excluding Cornutiplusia and Anagrapha) is a large 
genus of about 35 species. Its members are generally characterized by spined hind (and occasionally other) 
tibiae but this character state also occurs in several other plusiine taxa, particularly if occasional spines are 
counted (see Character analysis). The three taxa examined in this study also shared several other features 
of interest . They are the only members of the Plusiini to have a straight uncus (97) and a sclerotized , sharply 
pointed, hook-like harpe (149). Like Anagrapha, they have a subapical juxtal process (127) and a T8 cut 
back ventrally below the base of the anterior apophysis (189). This feature also occurs in the Autoplusiina 
(clade30). 

On Eichlin & Cunningham's (1978) phylogenetic tree, Syngrapha occupied the apex, as the sister-group 
of Plusia. They recognized a maximum of ten lineages within Syngrapha (excluding Anagrapha). I will 
discuss four of these in this section, following the topology of Eichlin & Cunningham's tree. S. surena 
(Grote) will be discussed under Palaeographa; the celsa and rectangula species-groups under clade 45; and 
Palaeographa, Syngrapha (s.str.) and Caloplusia in their own sections. More detailed descriptions of the 
Nearctic species are given by McDunnough (1944) and Eichlin & Cunningham (1978). 

5. ottolenguii was considered by Eichlin & Cunningham to be the most primitive Syngrapha. In many 
ways, the species is halfway between this genus and Autographa. S. ottolenguii has the elongate, heavily 
sclerotized antrum, the ventral edge of which projects beyond the ostium bursae (191; a feature seen in 
Palaeographa and Caloplusia but not in any Autographa) and also spined hind tibiae (Eichlin & 
Cunningham, 1978). However, the male genitalia are very similar to those of many Autographa species, 
especially in having a long clavus (all other syngraphas have reduced clavi). Given that the form of the 
female genitalia is likely to be apomorphic, while that of the male plesiomorphic, I will retain ottolenguii in 
Syngrapha. McDunnough (1944) and Franclemont & Todd (1983) took a different view and placed the 
species in Autographa. Either way, S. ottolenguii is clearly another 'critical species' and knowledge of its 
immature stages is imperative. The species is restricted to Alaska and the Yukon Territory, west of the 
Mackenzie River (Eichlin & Cunningham, 1978). 

The next group proposed by Eichlin & Cunningham was a paraphyletic assemblage of all the remaining 
syngraphas that lacked seta SV2 on larval segments Til and Till. Three lineages (excluding Anagrapha) 
were recognized within this group. The most primitive, in that its members retained the raduloid on the 
larval hypopharynx, comprised four species: S. selecta (Walker), S. epigaea, S. altera (Ottolengui) and S. 
octoscripta (Grote). The larvae of S. epigaea and S. selecta have been described by Eichlin & Cunningham 
(1978). The former appears to be highly polyphagous on various trees and shrubs, while records for 5. 
selecta require confirmation due to confusion with the superficially very similar S. viridisigma (q.v.). Like 
that species, S. selecta may also be shown to be restricted to conifers. S. octoscripta has been recorded as 
feeding on Vaccinium. This group of syngraphas occurs across the northern U.S.A. and southern Canada. 
S. altera is restricted to east of the Great Lakes; S. epigaea extends southward into Oregon, Wyoming and 
Colorado; 5. octoscripta ranges farther north to Alaska and Newfoundland; while S. selecta is so far known 
reliably from scattered localities (Eichlin & Cunningham, 1978). 

Of those species that lack both SV2 and Til and Till and a raduloid, S. viridisigma was considered the 
most primitive, due to genitalic and wing pattern similarities with the selecta species-group. As stated 
above, this species is frequently confused with the rarer S. selecta but the genitalia of the two species are 
distinct . The larva was described by Eichlin & Cunningham (1978) and various conifers were listed as larval 
foodplants. Some of these may, however, refer to S. selecta. S. viridisigma occurs throughout Canada, and 
is found also in the far north-eastern U.S.A. and in the Rocky Mountains as far south as New Mexico. 

The most advanced group of syngraphas lacking seta SV2 on Til and Till (excluding Anagrapha) was 
considered to be the diasema species-group. All these species have pale yellow hindwings, similarly 
patterned forewings and show similarities in genitalic structure (Eichlin & Cunningham, 1978). The 
included species were: 5. diasema, S. snowii, S. sackenii, S. borea, S. diversigna and S. orophila. The larval 
foodplants of only two species are known; Vaccinium (S. orophila; Crumb, 1956) and Betula and Populus 



SYSTEMATICS OF THE PLUSIINAE 165 

(5. diasema; Kostrowicki, 1961). S. diasema occurs throughout central Canada and southern Alaska 
(Eichlin & Cunningham, 1978), and is also found in Scandinavia and the north-eastern U.S.S.R. It is 
probably circumpolar in distribution (Kostrowicki, 1961). S. snowii is apparently endemic to New Mexico, 
while S. borea is found in the far north of Canada and in Greenland. S. diversigna occurs in the Canadian 
Rockies and southern Alaska, and is broadly sympatric with S. orophila, which extends further south to 
Wyoming. S. sackenii is found on the eastern slopes of the Rocky Mountains in the U.S.A. and Alberta, 
and appears to be sympatric with 5. orophila in that part of the latter's range in which S. diversigna is absent 
(Eichlin & Cunningham, 1978). 

Palaeographa [6(1)]. When Palaeographa was first described as a subgenus of Syngrapha (Kljutshko, 
1983), four species were included: S. interrogationis, S. alpina Ichinose, S. gilarovi Kljutshko and S. 
transbaikalensis (Staudinger). S. alpina had previously been synonymized with S. nyiwonis Matsumura 
(Sugi, 1982), and this latter taxon is also referable to Palaeographa. Other candidates include S. pyrenaica 
Hampson, 5. flammifera Chou & Lu and S. u-aureum (Guenee). These last three species have all, at one 
time or another, been considered synonyms of 5. interrogationis. S. transbaikalensis apparently grades into 
S. interrogationis in Korea (Ronkay, 1982) and may thus be yet another form of the latter. 

As for the previous major Syngrapha subgroup, those species that possess seta SV2 on segments Til and 
Till were also subdivided by Eichlin & Cunningham (1978) based upon the presence or absence of the 
raduloid. 5. interrogationis and 5. u-aureum (together with S. surena, see below) constituted that lineage in 
which the raduloid was retained and was therefore considered primitive. This taxon may correspond to 
Palaeographa but confirmation of the presence of the raduloid in the larvae of the other species is required. 
Also, at present, well-documented synapomorphies for the members of Palaeographa are needed to 
corroborate the monophyly of the subgenus. 

S. interrogationis (Fig. 93) is a widespread Holarctic species. It occurs in northern Canada and Alaska 
(Eichlin & Cunningham, 1978), throughout northern and western Europe (the British Isles, Iceland, 
Scandinavia) and east through central U.S.S.R. to the Pacific seaboard. S. pyrenaica is endemic to the 
Pyrenees and may be no more than a local race of 5. interrogationis. S. transbaikalensis, as its name 
suggests, occurs in the north-eastern Palaearctic region, east of Lake Baikal to Japan and Kamtchatka. S. 
nyiwonis is found in Japan and the neighbouring land areas of Sakhalin and the Kurile Islands (Ichinose, 
19636). Further details of the distribution and ecology of 'S. alpina' were provided by Kljutshko (1983), 
who also stated that S. gilarovi was restricted to the area of Kamtchatka. S. flammifera was described as an 
aberration of S. interrogationis by Hoyningen-Heune (1901) but was subsequently elevated to full species 
level, from subspecies (sic), by Chou & Lu (19796). Given that the collection locality of the original 
specimen of flammifera was Estonia and Chou & Lu presumably examined specimen(s) from China, then 
either S. flammifera is a very widespread (or disjunct) but rare and highly undercollected species or, as is 
generally considered, is merely a form of S. interrogationis. The diagnostic characters used by Chou & Lu 
(19796) ('transverse cordate stigma of [the] fore-wing and [the] truncate apex of the clavus') suggest that 
the latter hypothesis is most likely correct. S. u-aureum occurs throughout Canada and the north-eastern 
U.S.A., with an isolated record from Colorado (Eichlin & Cunningham, 1978). 

The life history of only one species is known, European S. interrogationis (Bretherton etal. , 1983). These 
authors give Calluna and Vaccinium (Ericaceae) as larval foodplants, while Kostrowicki (1961) added 
Erica and Urtica. S. u-aureum is said to feed on Vaccinium (Forbes, 1954) but no further details of its 
biology are known. 

A second species, S. surena, was associated with S. interrogationis by Eichlin & Cunningham (1978) on 
the basis of similar genitalia. S. surena is very rare, Hampson (1913) never saw it and Eichlin & 
Cunningham (1978) knew of less than a dozen specimens. It occurs in Newfoundland, Ontario, Quebec 
and Maine. 

Clade 45 [7(1), 24(1), 127(3)]. Three synapomorphic character states unite Syngrapha (s.str.) and 
Caloplusia as sister-groups. All members of both taxa (see below) have yellow hindwings (7) except S. 
parilis, in which they are white. A juxta with a subapical, sharp spine (127) is also almost universal but is 
absent in S. montana. Small eyes with a prominent rim (24) are restricted to a subset of potential members 
of clade 45; Caloplusia, Syngrapha devergens and S. ignea. All three of these conditions are absent in the 
putatively related celsa and rectangula species-groups, which I will now discuss, separately from Syngrapha 
(s.str.) and Caloplusia. 

Eichlin & Cunningham (1978) considered as most highly evolved, that branch of Syngrapha in which seta 
SV2 was present on segments Til and Till of the larva but the raduloid had been lost. Four lineages were 
recognized within this group. Caloplusia (q.v.), with spines on all the tibiae, was interpreted as the first 
lineage to diverge. Next was the rectangula species-group, which was considered to be closely related based 



166 IAN J. KITCHING 

on similarities in larval and genitalic structure. The celsa species-group formed the sister-group of 
Syngrapha (s.str.). Thus clade 45 was treated by Eichlin & Cunningham (1978) as polyphyletic (Farris, 
1974), and the three synapomorphies listed above would have to be interpreted as convergent develop- 
ments. Further study of Syngrapha is required to clarify this issue. 

The celsa species-group comprises only 5. celsa and S. angulidens . Both species feed as larvae on various 
conifers (Crumb, 1956; Ross& Evans, 1957; Prentice, 1962). Nothing further is known of the biology of 5. 
angulidens but the larva of 5. celsa was described by Crumb (1956) and Eichlin & Cunningham (1978). 5. 
celsa is restricted to the western slopes of the Rocky Mountains from Alaska to southern California, while 
S. angulidens is found on the eastern slopes, from southern Canada to Arizona. There are also records of 5. 
angulidens from New Hampshire and Nova Scotia in the east (Eichlin & Cunningham, 1978). 

The rectangula species-group comprises four species: S. rectangula, S. alias (Ottolengui), S. abstrusa 
Eichlin & Cunningham and S. cryptica Eichlin & Cunningham. The latter two species have only recently 
been separated from S. alias (Eichlin & Cunningham, 1978). Nothing is known about their biology but 
some of the records reported for S. alias may well apply to these species. Like the celsa-group, S. rectangula 
and 5. alias feed as larvae on diverse species of conifers (Crumb, 1956; Ross & Evans, 1957; Prentice, 1962; 
Tietz, 1972). This habit was, together with the presence of SV2 on Til and Till, sufficient for Crumb (1956) 
to consider proposing a separate genus (genus no. 11). However, the setal character is now known to be 
much more widespread and so only monophagy on conifers remains as a putative synapomorphy for the 
rectangula and celsa species-groups. 5. alias and 5. rectangula are found across the northern U.S.A. and 
southern Canada, although the former also extends along the Rocky Mountains north to the Yukon 
Territory and south to Arizona. The distributions of the other two species are likely to be incomplete, due 
to confusion with S. alias. S. cryptica has so far been recorded from Wisconsin, Michigan and New 
Hampshire, and S. abstrusa additionally from Ontario and Maine (Eichlin & Cunningham, 1978). 

Syngrapha (s.str.) [83(0)]/ Caloplusia [59(1), 63 (2), 64(2)]. I will deal with these two taxa together because 
the distinction between them has been blurred in recent years. Caloplusia was originally erected by Smith 
(1884) for those northern species of 'Plusia' (s.l.) with ovate eyes and spines on all the tibiae. However, 
there were no explicitly included species until C. hochenwarthi and 'C. devergens' were placed in the genus 
some seven years later (Smith, 1891). Warren (1913) designated C. hochenwarthi as type species in April 
1913 (Nye, 1975) and added C. tibetana (Staudinger) and C. composita Warren. Hampson (1913; 
November) subsequently employed the same type species and added 'C. virgula' (now Rachiplusia), C. 
alticola (Walker) and C. ignea (Grote). In both works, Caloplusia was distinguished from Syngrapha by the 
possession of spines on all the tibiae, rather than on just the hind tibiae (characters 63 and 64). Both authors 
also incorrectly cited Syngrapha ain (Hochenwarth) as the type species of that genus and thus S. devergens 
could be comfortably accommodated in Caloplusia. 

This error was corrected by McDunnough (1944) and devergens (together with alticola) was transferred 
to Syngrapha. McDunnough's discovery that neither of these two species had the extensive tibial spining 
claimed for them (an observation confirmed in the present study) allowed him to leave Caloplusia as a 
distinct genus. The separation was retained by Forbes (1954), Crumb (1956) and Kostrowicki (1961). The 
last author committed the same error as Hampson (1913) but on this occasion it was S. interrogationis that 
was mistakenly treated as the type species of Syngrapha. 

Eichlin & Cunningham (1978) held the opinion that the degree of tibial spining was not a character of 
genus-level significance and synonymized Caloplusia with Syngrapha. However, only ignea was explicitly 
transferred; hochenwarthi was included only by default. Franclemont & Todd (1983) followed this 
synonymy and also sank the Nearctic 5. alticola to the Palaearctic S. devergens. 

There are thus eight species in clade 45 as constituted in this study. For convenience, I shall retain the 
names Caloplusia and Syngrapha (s.str.). The latter includes S. devergens (Fig. 94), S. montana, S. 
microgamma, S. parilis (Fig. 96) (following Eichlin & Cunningham, 1978) and S. ain. The last species has 
never previously had its relationships within Syngrapha analysed. An examination of the male genitalia of 
S. ain figured by Ichinose (19626) showed that it has the same general facies and large, basal, strongly 
curved cornutus on the vesica as S. microgamma, and it would seem that the two may be sister-species. 
Caloplusia comprises C. hochenwarthi (Fig. 95), C. composita and C. ignea. 

The biology of none of the Nearctic species of Caloplusia or Syngrapha (s.str.) is known, nor of the 
Nearctic populations of the Holarctic species. In Europe, S. ain is recorded as feeding as a larva on Larix 
(Pinaceae); S. microgamma on Salix (Salicaceae); 5. devergens on Silene (Caryophyllaceae), Viola 
(Violaceae), Plantago (Plantaginaceae) and Geum (Rosaceae); and C. hochenwarthi on various umbellif- 
ers (Kostrowicki, 1961). Tietz (1972) added numerous arctic-alpine plants to the list for C. hochenwarthi 
(Antennaria, Carex, Castilleja, Gentiana, Penstemon, Salix and Vaccinium). The larva of C. hochenwarthi 
has been briefly described by Kirby (1889) , but the chaetotaxy of all the species remains virtually unknown. 






SYSTEMATICS OF THE PLUSIINAE 167 

A larva of S. ain examined in the course of this study proved to possess seta SV2 on segments Til and Till . 

S. montana is the only wholly Nearctic species of Syngrapha (s. str. ) . It occurs mostly in the north-eastern 
corner of the U.S.A. and adjacent areas of Canada south of the St Lawrence river. There are isolated 
records from Wisconsin and Saskatchewan. The other endemic Nearctic species is Caloplusia ignea, found 
mainly in the Rocky Mountains from Alaska to California and Colorado, with additional records across 
northern Canada (Eichlin & Cunningham, 1978). C. composita is known only from the mountains 
south-east of Lake Balkhash (U.S.S.R.), while C. hochenwarthi has a disjunct distribution in Scandinavia, 
the European Alps, Macedonia, and various localities throughout the U.S.S.R. from Armenia and the 
Caucasus to the Amur river and Kamtchatka (Kostrowicki, 1961). Syngrapha ain occurs in the mountains 
of central Europe and in the central U.S.S.R. from Lake Balkhash to the River Lena. There are also 
records from the Amur river (Kostrowicki, 1961). 

The remaining three species are Holarctic. 5. microgamma is present throughout central Europe, 
Scandinavia and along the Baltic coastline, with scattered records in the central U.S.S.R. and Kamtchatka, 
and throughout Canada and the northern U.S.A. , with an isolated record from Colorado. 5. devergens also 
occurs in the European Alps, Scandinavia and south-east of Lake Balkhash and in the Nearctic, extends 
throughout the Rocky Mountains from Alaska to Colorado, and across to Hudson Bay. 5. parilis is 
circumpolar at high latitudes in Canada, Alaska, Greenland, Scandinavia, the Arctic Ocean coasts and 
Novaja Zemlya (U.S.S.R.) (Eichlin & Cunningham, 1978; Kostrowicki, 1961). Its absence from localities 
further east is almost certainly due to lack of collecting in these areas. 

Intra-outgroup analysis 

The cladistic relationships between the outgroup taxa are not of primary importance in this study. 
However, several interesting patterns were found and it might be useful, for future studies, to discuss them 
briefly. I will only deal with six of the possible ten outgroup clades. 

The complement of clade 51 [38(0), 40(0), 48(1), 56(1), 178(1)]. If the cladogram shown in Fig. 27 is 
rerooted on the branch subtending clade 51, then a new clade is formed (the complement of 51) that 
hypothesises that the sister-group of the plusiines is clade 47 (Magusa + the stictopterines + the 
euteliines). As might be expected, given that this topology is only one of several possible (see 648-step 
cladograms), the data supporting this sister-group relationship are relatively weak. The Plusiinae and the 
members of clade 47 are unique only in possessing a broad marginal flange to the dorsal tegular arm (56). 
However, as noted earlier, the states of this character are not clear cut and may be subject to 
reinterpretation. A better character is the possession of a pleural pouch (178), although this must be 
interpreted as having been lost in Magusa. 

Thus, although clade 47 is a reasonable candidate for the sister-group of the Plusiinae , the evidence is far 
from unequivocal. Further analysis is necessary, including a detailed study of the relationships of the taxa 
comprising clade 47, before the sister-group of the Plusiinae can be named with any degree of certainty. 

Clade 47 [20(2), 22(0), 23(1), 26(1), 34(1), 37(1), 81(0), 82(1), 83(3), 107(1), 118(0), 156(1), 210(1)]. There 
are seven good synapomorphies to unite the Stictopterinae, Euteliinae and Magusa into a monophyletic 
group: scale bases on the wings arranged into regular transverse lines (20); bulbous eyes and a prominent 
frons (23, 26); the setae on the proboscis continuing almost to the apex (34); a smooth ventral edge to the 
basal two segments of the labial palps (37); and the form of the hair pencils (81) and St8 (82). In addition, 
the apex of the pleurite is recurved over the tegumen arm to articulate in a pocket on the inner edge (118; 
also seen in Calophasia); the aedeagus is very small compared to the rest of the genitalia (156; reversed in 
Paectes); and there are regular ridges on the corpus bursae (210; present also in Calophasia and Cucullia). 
However, the monophyly of clade 47 should not be accepted uncritically, given the paucity of included 
genera examined. Although Magusa and Stictoptera look very alike superficially, experience of similar 
situations in the Plusiinae (e.g. Anagrapha and Macdunnoughia confusa) suggests that this may be a poor 
guide to close cladistic relatedness. The genitalia of Stictoptera and Magusa in both sexes are very different. 
Also, Magusa is a trifine, with rudiments of a trifine hair pencil (75), whereas the stictopterines and 
euteliines are unequivocally quadrifine with no trace of trifine hair pencil components. Although these 
characters may be prone to frequent reversal and convergence, they do suggest that the support for clade 47 
may not be as convincing as it may first appear. Also, the sternal hair pencils of Magusa are markedly more 
complex than any seen in the Stictopterinae and Euteliinae, and studies to confirm the homologies 
proposed in the current investigation are required. Thus, while there can be little doubt that the 
Stictopterinae and Euteliinae are sister-taxa (see below), the conclusion that Magusa is their sister-group 
must still be considered open. 



168 IAN J. KITCHING 

Clade 48 [13(0), 14(3), 15(0), 17(1), 18(0), 25(1), 43(1), 52(2), 57(1), 62(1), 66(1), 68(1), 70(1), 72(1), 
73(3), 114(1), 123(1), 153(1), 161(1), 179(1), 180(1), 190(1), 195(1), 202(1), 211(1)]. Next to the 
Abrostolini, clade 48, the Euteliinae + Stictopterinae, is the best corroborated clade found in this study. 
There are no fewer than 11 unique and unreversed synapomorphies to support it, together with a further 
nine in which the apomorphic state has been subsequently lost in one of the included genera and another 
four where a convergent development has been recorded in only a single genus outside the clade. 

The euteliines and stictopterines share the following good synapomorphies: a reduced female frenulum 
(14), a full quadrifine hindwing venation (15; probably synapomorphic only in the context of the taxa 
included in this study , but , given Magusa is trifine , maybe a reversal) ; a frons scaled only at the edge (25) ; a 
patagium with a drawn-out attachment point (57); weak SV and V hind basitarsal spines (62); hindlegs 
shorter than the midlegs (66); dorsal furcal arms that are curved posteriorly (70); the form of the internal 
flange on T2 (73); ventrally or subventrally positioned valves (153); a small, oval plate in the ductus 
ejaculatoris (161); and anal papillae in which the inner surfaces are turned to face at least partially 
posteriorly (179). In addition, the Stictopterinae show partially-developed 'wings' on the proboscis 
styloconic sensilla (33) (Fig. 113). If this condition is interpreted as intermediate between the plesiomor- 
phic state 1 (Fig. 112) and the apomorphic state 2 (Figs 114-116), then it further corroborates the 
monophyly of both clade 48 and the Euteliinae. 

They also share the following putative synapomorphies but these are either lost in one of the constituent 
taxa (indicated by '— genus' in parentheses) or occur in another genus outside clade 48 ('+ genus' in 
parentheses) . The character states involved are : underside of the forewing cell without setose scales ( 13 , + 
Chalcopasta); forewing vein R2+R3+R4 markedly converging on R^ then curving away distally (17; - 
Paectes) ; vom Rath's organ conspicuously papillate (43; - Anuga); ventral arm of the tegula spatulate (52; 
— Stictoptera); hindleg inner apical spurs approximately three-quarters length of basitarsus (68; - 
Lophoptera); abdomen distinctly tapered posteriorly (72; - Lophoptera); dorsal and ventral flanges of 
vinculum arms fused across to give a flattened, cone-shaped structure (114; -I- Rachiplusia); vinculum and 
tegumen broadly joined, pleurite completely fused to tegumen (123; + Diloba); inner surface of the anal 
papillae densely setose (180; + Calophasia); anterior apophyses absent (190; — Stictoptera); female T8 
ventrally produced anteriorly to lie alongside antrum dorsally and exceeding it ( 195 ; - Stictoptera) ; female 
T8 represented by a thin band dorsally (202; - Stictoptera); and the corpus bursae with an invaginated 
signum (211; — Anuga). 

Examination of the preceding 13 characters reveals that there is no strong pattern to the homoplasies 
that would imply that either subfamily is paraphyletic relative to the other. It could be argued that 
Lophoptera is more closely related to the euteliines than it is to Stictoptera, because it shares four putative 
synapomorphic features missing in Stictoptera (52, 190, 195, 202). However, there are two character states 
(68, 72) that imply the opposite relationship and, given that there are other characters to corroborate the 
monophyly of the two subfamilies, the six states just mentioned must be interpreted as homoplasies. 
Certainly, no firm conclusions should be drawn about the internal structure of clade 48 when only 4 of the 
40 or so generic-level taxa have been examined, but equally, I think it is unlikely that the monophyly of the 
Stictopterinae and of the Euteliinae will be refuted. 

Although Hampson (1912) recognized the reduced female frenulum in the Stictopterinae and Eute- 
liinae, he misinterpreted its structure in the latter group, referring to it as being 'single'. He considered the 
two subfamilies to be parallel developments from the 'Ophiderinae'. Seitz (1937) also observed that the 
euteliines and stictopterines had a number of features in common, including the large, bulbous eyes (23), 
from which he concluded that they were closely related. 

The recognition of a close relationship suffered a setback with Richards' (1932) study of the tympanal 
organs of noctuids. He observed a secondary tympanal hood (pleural pouch) in the stictopterines and the 
plusiines and derived the latter from the former. The euteliines were thought to have much in common with 
the acontiines and although a membranous secondary tympanal hood was observed in Eutelia, the 
connection with the Stictopterinae (and Plusiinae) was not made. Forbes (1954) referred only to the 
stictopterine-plusiine relationship, suggesting that these two subfamilies might perhaps be combined, and 
the euteliine-stictopterine connection was lost. 

Recently, in a study of several Bornean noctuid subfamilies, Holloway (1985) resurrected the hypothesis 
of a euteliine-stictopterine connection when he noted the presence of a small, oval sclerotization in the 
ductus ejaculatoris (character 161) of the members of these two subfamilies. He also noted several other 
interesting similarities (e.g. the morphology of the anal papillae) and these formed the basis of the 
characters employed in this study. 

Clade 49: the Stictopterinae [8(1), 14(5), 61(1), 62(2), 121(1), 151(1), 181(1)]. Five unique and unreversed 
synapomorphies were found for the two stictopterine genera included in the data set. These are: a hyaline 



SYSTEMATICS OF THE PLUSIINAE 169 

hind wing base (8); a female frenulum reduced to a single seta (14); suppressed SV and V spines on the 
basal half of the hindbasitarsus(62); a deeply cleft valve (151); and apically 'frayed' anal papillae (181). Of 
these, not all stictopterines show the apomorphic states of characters 8 (e.g. Odontodes) and 151 (e.g. the 
Lophoptera khasiana Warren species-group) (Holloway, 1985). Character 62 requires confirmation from 
the other genera but Holloway (1985) found all Bornean stictopterines to have the apomorphic states of 
characters 14 and 181. To these he added three more putative synapomorphies: the form of the tympanal 
hood, the general wing facies and a distinctive Y-shaped pupal cremaster (Gardner, 19486). The 
monophyly of the subfamily thus seems reasonably well corroborated. 

Holloway also provided brief reviews of the morphology (both of adults and immatures), biology and 
zoogeography of the subfamily. In particular, he concluded that the two major lineages of stictopterines 
have different larval foodplant preferences. The Stictoptera genus-group appears to be restricted to 
members of the Clusiaceae (Guttiferae), while the Lophoptera genus-group is primarily recorded from the 
Dipterocarpaceae. According to Thome (1976) and Heywood (1978), these two families are closely 
related, both being members of the Theales, but more recent classifications of the angiosperms (Dahlgren, 
1983; Thome, 1983) disagree, placing the dipterocarps in the Malvales. Members of the Stictoptera 
genus-group may be of potential economic importance as defoliators of crop trees such as Garcinia 
(mangosteen) (Holloway, 1985). 

Clade50: the Euteliinae [33(2), 35(2), 36(1), 46(1), 47(0), 58(1), 68(2), 71(1), 74(1), 107(0), 115(1), 122(1), 
129(1), 148(0), 152(1), 157(1)]. The Euteliinae are even better defined and more autapomorphic than their 
sister-taxon, with no fewer than ten unique and unreversed synapomorphies to corroborate their 
monophyly. Only Paectes and Anuga were found to possess the following features: winged styloconic 
sensilla on the proboscis (33) (but see clade 48 above), which is nodular only at the extreme apex (35) and 
has regular, dense, basal rows of microspines (36); asymmetrical, bipectinate male antennae (46), with the 
pectinations bearing only a single, small, apical seta (47); hindleg inner apical spurs that equal or exceed 
the hind basitarsus in length (68); two internal flanges on St2 (74); vinculum arms that are distally parallel 
or convergent (115); socii (122); a crescent- or boomerang-shaped, weakly sclerotized juxta (129); and a 
somewhat trapezoid valve (152). 

Of these, however, the antennae are ciliate or filiform in Anigraea Walker and Targalla Walker 
respectively; the valves of other genera are more varied in shape; and socii are frequently absent 
(Holloway, 1985). Many of the other characters listed above require confirmation but all genera appear to 
possess the flanges on St2, which may be associated with the characteristic resting position of the adults 
(Holloway, 1985; Kitching, 1984). 

Reviews of the morphology of the adults and immatures, biology and zoogeography of the Euteliinae 
have been presented by Holloway (1985). The lack of a pupal cremaster and a bias towards the 
Anacardiaceae as larval foodplants (but see also Peigler, 1986) may also represent potential apomorphies 
for the subfamily. Several genera have been recorded as agricultural pests, especially of mango (Man- 
gifera). 

Holloway (1985) did not propose a cladistic hypothesis for the euteliine genera. Mell (1943) recognized 
four genus-groups based on Penicillaria Guenee (as Bombotelia Hampson), Eutelia Hiibner (as Phlo- 
gophora Treitschke), Anugal Mimanuga Warren and Paectes, but only examined a very small number of 
species. Thus, the internal relationships of the Euteliinae remain to be elucidated. 

Clade 55 [20(0), 21(1), 31(0), 32(2), 34(2), 46(2), 55(1), 71(1), 96(1), 208(1)]. The position of the enigmatic 
genus Diloba, and its single species, D. caeruleocephala, has changed with almost monotonous regularity. I 
gave a brief historical review (Kitching, 1984), charting its course from the Bombyces, via the Noctuidae: 
Bombyciformes, Notodontidae: Dilobinae, Noctuidae: Plusiinae, Thyatiridae, Noctuidae: Acronictinae, 
Noctuidae: Pantheinae, Lymantriidae and Dilobidae, to the Noctuidae: Cuculliinae. Diloba has certainly 
been around! 

Possible inclusion in the Cuculliinae was first mooted by Tarns (quoted in Varley, 1962), who suggested a 
relationship with the American genus Psaphida Walker. Minet (1982), in a critique of Kiriakoff s (1970) 
work on the tympanal organ of Diloba, independently hypothesised a place for the genus in the 
Cuculliinae, associating it with Episema Ochsenheimer. In this study, I decided to follow up Tarns' 
suggestion to test the relationships of Diloba. The members of the cuculliine tribe Psaphidini are mostly 
Nearctic and poorly represented in the BMNH collection. However, specimens of the Holarctic genus 
Brachionycha are more numerous, and this taxon was chosen to represent the Psaphidini, although there 
have been reservations concerning its placement in this tribe (Forbes, 1954). 

I found six putative synapomorphies that were unique to Brachionycha and Diloba. These would appear 
to confirm the place of the latter in both the Noctuidae and the Cuculliinae. However, a more extensive 



170 IAN J. KITCHING 

investigation is required to confirm the relationship of these two genera with the Psaphidini and/or 
Episema. The synapomorphies are: the presence of a vestigial remnant of forewing vein CuP (21); a 
reduced, non-functional proboscis with no distal styloconic sensilla and setae continuing to the apex (32, 
31, 34); the ventral arm of the tegula produced on a lateral extension of the main body (55); and an 
apparently small corpus bursae, but with extremely elastic walls (208). 

Some of thse conditions are somewhat unconvincing as synapomorphies (especially the reduction 
characters) but the unusual tegulae have not so far been found elsewhere. There are numerous differences 
between the two genera but it was not expected that Brachionycha would prove to be the sister-group of 
Diloba. However, I hope that I have shown that future studies of the relationships of Diloba should 
concentrate on the Psaphidini and not on the supposed notodontid connection. 

Consensus classification 

From the preceding evaluation of the preferred 648-step cladogram, it is clear that there are both clades 
and characters of highly dubious value. For example, the apomorphic state of character 59 has been gained 
and lost six times, while the monophyly of clade 38 is supported by only a single state change in a character 
(105) that has a very low consistency index (0-111). For this reason, I performed one final analysis of the 
data set and will base my classification of the Plusiinae on this, rather than on the preferred 648-step 
cladogram. 

Analytical procedure 

The data set was modified for re-analysis as follows. I considered the monophyly of the Plusiinae to have 
been reasonably well-demonstrated and thus decided to remove the 1 1 outgroup taxa from the matrix. This 
also allowed me to exclude 55 characters that exhibited state changes only among the outgroups. These 
were: 8, 13, 17, 21, 23, 25, 26, 27, 30, 34, 36, 37, 38, 40, 43, 44, 45, 48, 53, 54, 55, 56, 57, 62, 66, 67, 68, 69, 
70, 71, 72, 74, 75, 81, 82, 96, 115, 122, 123, 129, 135, 151, 152, 153, 156, 161, 179, 180, 181, 190, 195, 202, 
208, 210 and 211. A further five characters (20, 22, 46, 47, 114) were discarded because, after removal of 
the outgroups, they became autapomorphic for a terminal taxon within the Plusiinae. Finally, all 
characters that showed consistency indices of ^ 0-250 were deleted from the data set (5, 6, 12, 16, 18, 19, 
42, 58, 59, 105, 107, 110, 144, 162, 191, 196, 201, 205, 209), with the following additions and exceptions. 
Characters 65 (c = 0-750) and 83 (c = 0-313) were also removed. There were doubts regarding the 
distinction of the states of the first; while the second (male sternal hair pencil development) showed 
substantial homoplasy, and its relatively large consistency index is due, in part, to the large number of 
states (5) scored. Ten characters were also left in the matrix despite having consistency indices of =£ 0-250: 
3, 41, 97, 108, 131, 141, 148, 149, 189, and 214. These were generally characters that exhibited homoplasy 
only within the Plusiinae. Their apomorphic conditions also helped define high level clades in the preferred 
648-step cladogram and might, therefore, be expected to exert a significant influence in a re-analysis. For 
example, 141 appears to be potentially important in the Autoplusiina, while 189 concerns the possible 
relationship between Syngrapha (s.\.),Anagrapha and clade 30. 

As before, Shensiplusia was also omitted from the analysis. 

Finally, the removal of the outgroups allowed character 14 to be analysed as an ordered character; all 
other unordered characters were treated as before. 

The analysis was performed using the PAUP options HOLD = 3, SWAP = GLOBAL, MULPARS, 
NORELEASE; all other options being set to their default values. It was hoped that the deletion of about 
one-fifth of the taxa and over a third of the characters would speed up the computations. However, the 
opposite result was obtained; the analysis slowed considerably and took an almost excessive amount of 
computing time. I would therefore suggest that if anyone intends to delete such a large amount of data 'in 
program', as was done here, then they circumvent the tree-building routine, which took most of the time, 
by entering a USER-TREE that is expected to approximate the end result and perform branch-swapping 
on that topology. An even better approach would probably be to recast the data physically to remove the 
required taxa and characters and analyse this smaller data matrix. Either of these techniques should 
decrease the run times of large reduced data sets significantly. 

Results 

As might have been expected, the MAXTREE default maximum of 50 cladograms was obtained. They 
were 301 steps in length and had consistency indices of 0-584. Two observations can be made from these 
figures. First, exclusion of the outgroups and characters of low consistency eliminated over half the steps of 
the earlier cladograms (i.e. 346 of the original 647) . However , the consistency index did not rise by as much 



SYSTEMATICS OF THE PLUSIINAE 171 

as perhaps might have been expected, 0-475 to 0-584. This is primarily because the rise in consistency due to 
the loss of the low consistency characters was counterbalanced to some degree by a lowering in consistency 
due to the loss of many high consistency characters concerned with the outgroup interrelationships. 

Examination of the 50 cladograms revealed a series of alternative subtopologies similar, but not 
identical, to those obtained for the 648-step cladograms. (The clade numbers used subsequently are those 
employed on the preferred cladogram and imply the same constituent taxa, though not necessarily the 
same topology.) 

First, the positions of Agrapha and Plusiopalpa varied. Three alternative topologies were recorded. On 
the first, Plusiopalpa is placed as the sister-group of Argyrogramma + Stigmoplusia, with Agrapha as the 
sister-group of Ctenoplusia + Acanthoplusia. The other two topologies are the same as those found for 
clade 14 on the 648-step cladograms (see above). 

Dactyloplusia has moved to become the sister-group of Thysanoplusia, rather than, as before, forming 
the sister-group of Chrysodeixis + Pseudoplusia. This new clade is placed either as the sister-group of 
Trichoplusia or as the sister-group of Trichoplusia + clade 12. Otherwise, the Argyrogrammatini display 
the same topology as the preferred 648-state cladogram. 

In the Euchalciina, there is ambiguity as to the position of Panchrysia + Chrysanympha. One 
subtopology places this clade as the sister-group of Euchalcia + Adeva, while the other treats it as the 
sister-group of clade 24, as on the preferred cladogram. Otherwise the Euchalciina is unchanged. 

The exclusion of the characters of low consistency, as might have been expected, had a significant effect 
upon the topology of the Autoplusiina and Plusiina. Clade 30 is recovered by and large intact, although 
Loboplusia is placed as the sister-group of clade 32 + 35, with Erythroplasia as the sister-group to this 
clade. However, Loboplusia and Erythroplasia can exchange positions. 

At the base of the Autoplusiina, and within the Plusiina, there is what can only be described as near 
chaos, with at least 13 alternative subtopologies being found. Syngrapha (s.l.) is recovered intact as before. 
A new clade is also consistently obtained, comprising two pairs of taxa in a sister-group relationship; 
Lophoplusia + Plusia and Autographa bractea + Cornutiplusia. However, the relationships of these, 
Autographa gamma, clade 30, Anagrapha, Rachiplusia and Diachrysia appear to be able to take on any 
form. For example, at one extreme, A nagrapha forms the sister-group of Rachiplusia , which together form 
the sister-group of the rest of the Autoplusiina, while Diachrysia forms the sister-group of clade 39. At the 
other extreme, clade 37 is recovered intact but Rachiplusia is interpreted as the sister-group of clade 30 + 
37. Numerous other alternative subtopologies were found, most of which comprise variations on the 
placements of Anagrapha, Rachiplusia and Diachrysia. Such a result was not unexpected in view of the 
evaluation of these taxa performed above and the characters deleted in the present analysis. 

Along the main stem of the cladograms, no change in the topology was expected nor found. The cladistic 
relationships among the tribes and subtribes is identical to that of the preferred 648-step cladogram, 
namely: (Omorphinini (Abrostolini (Euchalciina (Autoplusiina + Plusiina)))). 

To extract the maximally predictive classification of the Plusiinae from these alternatives, a manual strict 
(Nelson) consensus cladogram analysis (Nelson, 1979) was performed. The ambiguity in clade 14 is 
reduced to a terminal tetrachotomy between Plusiopalpa, Agrapha, Argyrogramma + Stigmoplusia and 
Ctenoplusia + Acanthoplusia. Similarly, trichotomous relationships are assumed between clade 12, 
Trichoplusia and Thysanoplusia + Dactyloplusia; and between clades 23, 25 and 26 in the Euchalciina. 
Rachiplusia is removed from the Autoplusiina and placed with Anagrapha and Diachrysia in a pentachoto- 
mous relationship with clades 30 and 37. This best reflects the uncertainty of relationship of the three single 
genera concerned. Within clade 30, Loboplusia forms a trichotomy with Erythroplusia and clade 32 + 35; 
while in clade 37, Autographa gamma, Syngrapha (s.l.) and Lophoplusia/ Plusial Autographa bractea/ 
Cornutiplusia are also trichotomously related. 

Finally , in recognition of the fact that , on the preferred 648-step cladogram , neither clade 12 nor clade 14 
was supported by good synapomorphies, and that, for example, blade-like valve setae (133) may 
eventually prove to be a good synapomorphy for clades 13 and 16, the branches subtending clades 12 and 14 
were collapsed, leaving a terminal heptachotomy within the Argyrogrammatini. This consensus cladogram 
(with the addition of Pseudochalcia, Desertoplusia and Shensiplusia; see below) is shown in Figs 29, 30, and 
will form the basis of the formal higher classification of the Plusiinae presented in this study. 

The positions of Shensiplusia, Desertoplusia and Pseudochalcia 

Shensiplusia. Shensiplusia was described by Chou & Lu (1974) to accommodate the species 5. nigribursa, 
known only from the holotype female. It was said to resemble Argyrogramma (in the broadest sense) and, 
specifically, to be very similar in external appearance to Acanthoplusia (as Argyrogramma) agnata. S. 
nigribursa differed in being larger and in having more metallic iridescent scaling on the forewings. This 



172 IAN J. KITCHING 

PLUSIINI 

Argyrogramma 
Stigmoplusia 
Plusiopalpa 
Ctenoplusia 
Acanthoplusia 
# Agrapha 

Plusiotricha (Eutheiaplusia) 

Plusiotricha (Plusiotricha) 

Trichoplusia 

Thysanoplusia 

Dactyloplusia 

Zonoplusia 

Chrysodeixis (Chrysodeixis) 

Chrysodeixis (Pseudoplusia) 

Anadevidia 

Shensiplusia 

Abrostola 

Mouralia 

Omorphina 

Fig. 29 The consensus classification of the tribes Omorphinini, Abrostolini and Argyrogrammatini. The 
genus Shensiplusia is incertae sedis in the Argyrogrammatini. 




description, together with Chou & Lu's record for discovering 'new' species in China, suggests that S. 
nigribursa may eventually prove synonymous with A. agnata or a related species. However, the female 
genitalia of A. agnata figured by Ichinose (19626: pi. 9, fig. 6) do not correspond exactly to those of 5. 
nigribursa (Chou & Lu, 1974: fig. 16). In particular, A. agnata does not appear to have the corpus bursae 
covered internally with the numerous sclerotized spinules that give S. nigribursa its name. Chou & Lu 
(1974) also stated that the corpus bursae of S. nigribursa was somewhat similar to that of Chrysodeixis (as 
Neoplusia) acuta. 

Although Chou & Lu provided little information useful in placing Shensiplusia within the Plusiinae, it 
seems likely that the genus is an argyrogrammatine. However, further resolution of its position is 
impossible at present (in particular, in the absence of the male) and in recognition of this, I have placed 
Shensiplusia at the base of the Argyrogrammatini, in a trichotomy with Anadevidia and clade 6 (Fig. 29). 

Needless to say, nothing is known of the biology of Shensiplusia, except that it was taken at Wukung in 
Shensi Province, China. 

Desertoplusia. I learned of the descriptions of Desertoplusia and Pseudochalcia (Kljutshko, 1984) too late 
to include their type species, D. bella (Christoph) and P. inconspicua (Graeser), in the full analysis. I thus 
have to be content with placing them as best I can onto the cladogram (Fig. 30) constructed from the other 
genera. To this end, they will also provide useful tests of the robustness of the arrangement and its 
supporting characters. 

Desertoplusia contains two species, D. bella (Fig. 70) and D. paghmana (Wiltshire). In superficial 
appearance, they resemble miniature autographas and were placed in that genus by Wiltshire (1971) and as 
its sister-group by Kljutshko (19856). However, an examination of the male genitalia of both species, as 



SYSTEMATICS OF THE PLUSIINAE 



173 




Syngrapha (Caloplusia) 

Syngrapha (Syngrapha) 

Syngrapha (Palaeographa) 

Lophoplusia 

Plusia 

Cornutiplusia 

"Autographa" bractea 

Autographa gamma 

Anagrapha 

Diachrysia 

Rachiplusia 

Autoplusia 

Antoculeora 

Sclerogenia 

Allagrapha 

Macdunnoughia (Macdunnoughia) 

Macdunnoughia (Puriplusia) 

Loboplusia 

Erythroplasia 

Lamprotes 

Pseudeva 

Plusidia 

Desertoplusia 

Euchalcia (Euchalcia) 

Euchalcia (Adeva) 

Panchrysia 

Chrysanympha 

Polychrysia 

Eosphoropteryx 

Pseudochalcia 



Fig. 30 The consensus classification of the tribe Plusiini, comprising the subtribes Euchalciina, Autoplu- 
siina and Plusiina. The genus Pseudochalcia is incertae sedis in the Euchalciina, while the genera 
Rachiplusia, Diachrysia and Anagrapha are incertae sedis in the Autoplusiina + Plusiina. Loboplusia 
and Erythroplusia are incertae sedis in the Autoplusiina. 



174 IAN J. KITCHING 

figured by Wiltshire (1971), reveals that they have simple oval valves (147), reduced clavi (144:1) and 
subapical cornuti that arise from circular, concave plates (164). These character states indicate that 
Desertoplusia belongs respectively to clades 21, 24 and 25. However, Desertoplusia retains the plesiomor- 
phic plusiine forewing colour pattern and thus in Fig. 30, it is placed as the sister-group of Euchalcia + 
Adeva. However, it should be noted that future studies may position the genus internal to clade 25. 

D. bella is recorded in the BMNH collection only from Ashkhabad in Turkmeniya S.S.R. D. paghmana 
comprises two subspecies. D. p. paghmana is confined to the Paghman Mountains, north-west of Kabul in 
Afghanistan, while D. p. paupera is found only near Quetta, Pakistan (Wiltshire, 1971). Nothing is known 
about the biology of either species but they appear to be part of the same steppe-montane fauna as 
Euchalcia and Pseudochalcia. Being a euchalciine, the larval foodplant is predicted to be a member of the 
Ranunculaceae or, much less likely, the Boraginaceae. 



Pseudochalcia. I mentioned above in the clade analysis that Kljutshko (1984) had erected Pseudochalcia to 
accommodate P. inconspicua (Fig. 69), thereby formalizing Dufay's (1968) conclusion that the species did 
not belong in Euchalcia. Kljutshko (1985ft) interpreted Pseudochalcia to be the sister-group of Euchalcia, 
based on the shape of the juxta apex. In both genera, this forms a somewhat recurved, blunt lobe. In this 
study, however, a similar condition was found in Diachrysia chryson, Autographa gamma, Erythroplusia 
rutulifrons, Antoculeora ornatissima, Adeva albavitta and Chrysanympha formosa. Examination of the 
male and female genitalia figured by Kostrowicki (1961) yields little further data to help place Pseudochal- 
cia. The valve is roughly quadrate in shape and thus resembles certain Euchalcia species. However, the 
subapical cornutus is absent, the vesica ornamentation consisting primarily of a transverse row of straight 
cornuti about halfway(?) along the vesica. The sacculus is armed apically with two short, divergent, sharp 
points (saccular sheets?) and appears to have a basal, broad triangular point. These features suggest a 
possible relationship with certain autoplusiine taxa. However, the apex of the corpus bursae is membra- 
nous and lacks sclerotized ridges. Thus, like Dufay ( 1968) , I do not know what the detailed relationships of 
Pseudochalcia are. It is certainly a member of the Plusiini and possibly the Euchalciina, although inclusion 
in the Autoplusiina cannot be entirely ruled out. In recognition of this uncertainty, I have placed 
Pseudochalcia in a trichotomous relationship in the Euchalciina with Eosphoropteryx and clade 21 (Fig. 
30). 

Nothing is known of the biology of P. inconspicua. It is restricted to the mountains of Central Asia 
(Kostrowicki, 1961; Kljutshko, 1984). 



The higher classification of the Plusiinae 

The formal classification of the plusiine generic and higher level taxa given in Appendix 5 was constructed 
using the conventions for annotated Linnaean hierarchies defined and described by Wiley (1979; 1981). 
The hierarchy is fully sequenced (Nelson, 1974), that is, 'taxa forming an asymmetrical part of a phylogenetic 
tree may be placed at the same categorical rank and sequenced in phylogenetic order of origin' (Wiley, 
1981: 209). This has the effect of reducing the number of named subordinate taxa, the proliferation of 
which has long been a target for anti-cladists (e.g. Colless, 1977). I have reduced this number further by 
adopting informal genus-groups and, twice, genus-complexes. This approach was adopted because of the 
intrinsic uncertainty regarding several areas of the underlying cladogram. 

Taxa forming parts of polytomies 'are given equivalent ranks and placed sedis mutabilis at the level of the 
hierarchy at which their relationships to other taxa are known' (Wiley, 1981: 211). This convention 
differentiates those parts of the sequences that are dichotomous from those that are polytomous. It 
indicates that the taxa labelled sedis mutabilis are of interchangeable position. 

Finally, five genera were identified as being incertae sedis, of uncertain position. They are 'placed in the 
hierarchy ... at the level their relationships are best understood' (Wiley, 1981: 212). For example, 
Shensiplusia is considered to be an argyrogrammatine, but its sister-group could be the whole tribe or any 
clade within the tribe. It is thus placed incertae sedis at the level of the Argyrogrammatini. 

The application of these conventions allows the construction of a Linnaean hierarchy from which the 
structure of the underlying cladogram is immediately apparent. Further conventions were formulated by 
Wiley (1979; 1981) but were not necessary in the present context. 



SYSTEMATICS OF THE PLUSIINAE 175 

The Plusiinae are divided into four tribes: the monotypic Omorphinini Chou & Lu, the Abrostolini 
Eichlin & Cunningham, the Argyrogrammatini Eichlin & Cunningham and the Plusiini Boisduval. The 
Abrostolini comprises two genera, Abrostola and Mouralia. 

Within the Argyrogrammatini, as noted above, Shensiplusia is placed incertae sedis. The first level of 
sequencing comprises three genera (Anadevidia, Chrysodeixis and Zonophisia) and the terminal Argyrog- 
ramma-complex of genera and genus-groups. Chrysodeixis contains two subgenera, Chrysodeixis (s.str.) 
and Pseudoplusia stat. n. The Argyrogramma-comp\e\ consists of seven lineages, all placed sedis 
mutabilis. These consist of five genera (Trichoplusia, Plusiotricha, Ctenoplusia, Agrapha and Plusiopalpa) 
and two genus-groups. Genus-groups were erected generally where the evidence in favour of synonymy 
was considered inadequate. Thus Thysanoplusia stat. rev. and Dactyloplusia are placed together in the 
Thysanoplusia genus-group, and Argyrogramma and Stigmoplusia in the Argyrogramma genus-group. In 
contrast, Eutheiaplusia stat. n. and Acanthoplusia stat. rev. are reduced to subgeneric rank within 
Plusiotricha and Ctenoplusia respectively. This approach, rather than full synonymy, was adopted because 
I consider that both Eutheiaplusia and Acanthoplusia represent distinct monophyletic units within their 
respective genera (see Clade analysis). As concluded above (Clade analysis), and contrary to the 
(unstated) opinions of Franclemont & Todd (1983), I have found no convincing evidence for accepting a 
synonymy of Ctenoplusia with Agrapha. I thus reinstate Ctenoplusia stat. rev. as a genus separate from 
Agrapha. 

The Plusiini is divided into three subtribes: the Euchalciina Chou & Lu, the Autoplusiina subtrib. n. and 
the Plusiina Boisduval. 

Within the Euchalciina, Pseudochalcia is placed basally incertae sedis. The first level of sequencing 
comprises Eosphoropteryx, Polychrysia and the Panchrysia, Euchalcia and Lamprotes genus-groups. The 
last three of these are placed sedis mutabilis. They contain, respectively, Panchrysia and Chrysanympha, 
Desertoplusia and Euchalcia, and (sequenced) Plusidia, Lamprotes and Pseudeva. Following the sugges- 
tions of Kostrowicki (1961), Dufay, (1968) and Franclemont & Todd (1983), Adeva is placed within 
Euchalcia, but as the subgenus Euchalcia (Adeva) stat. n., until such time as the internal structure of 
Euchalcia (s.str.) is resolved. 

Although the taxon Autoplusiina + Plusiina is not formally named, three genera are placed incertae sedis 
at this level in the hierarchy. Rachiplusia will probably be shown eventually to belong to the Autoplusiina 
but I can only speculate that this may also be the correct tribe for Anagrapha and Diachrysia. 

Within the Autoplusiina, Erythroplusia and Loboplusia are placed basally incertae sedis . The remaining 
two genus-groups form a fully dichotomous and sequenced clade. The Autoplusia genus-group comprises 
Sclerogenia, Antoculeora and Autoplusia; the Macdunnoughia genus-group consists of Allagrapha and 
Macdunnoughia. The latter genus includes Puriplusia as a subgenus. 

Three groups are recognized within the Plusiina, in trichotomous relationship. That represented by 
Autographa gamma is probably paraphyletic and requires further work. Syngrapha (s.l.) is recognized as a 
monophyletic group comprising three sequenced subgenera (at least): Palaeographa, Syngrapha (s.str.) 
and Caloplusia stat. rev. The final taxon in this group is the P/us/a-complex, comprising two genus-groups 
based upon Plusia and Cornutiplusia stat. rev. The former also includes Lophoplusia stat. n., the latter, the 
group of Autographa represented by A. bractea. Given that there is at least one further lineage within that 
genus (those species possessing seta SV2 on segment Al of the larva), Autographa is at least triphyletic and 
ought eventually to be subdivided. Until a thorough investigation is undertaken, however, formal naming 
of the various lineages should be avoided. 



Comparison with previous classifications and phytogenies 

After Eichlin & Cunningham (1978) proposed their phylogeny of the Nearctic Plusiinae, they noted that 
although it and those of Kostrowicki (1961) and Ichinose (19626) had been 'derived independently . . . 
studying different faunal areas . . . the basic evolutionary lines are essentially the same'. Subsequently, 
two further dendrogrammatic representations of the higher classification of the Plusiinae were published 
(Chou& Lu, 1979c; Kljutshko, 1985a; b). A comparison of these five hierarchies with that proposed in the 
present study shows that Eichlin & Cunningham's observation still, by and large, holds true. However, 
there are discrepancies between each and my own conclusions that deserve brief discussion. 

Kostrowicki (1961) (Fig. 1 v Figs 29, 30). Kostrowicki's study was, by a narrow margin, the first new 
attempt to analyse the higher classification of the Plusiinae since Hampson (1913). It was based upon the 
Palaearctic fauna and thus included very few representatives of the primarily tropical Argyrogrammatini. 



176 IAN J. KITCHING 

If it is assumed that Kostrowicki would have converted his 'scheme of relationships' directly into a 
classification (which is unlikely but a necessary assumption if a meaningful comparison is to be made), then 
the following points of concordance and disagreement with my classification can be noted. 

Abrostola and Mouralia were considered to be the most primitive group of plusiines (stated implicitly). 
Omorphina was erroneously transferred, without explanation, to the Acontiinae (as Erastriinae). Within 
the Plusiinae examined, Kostrowicki's first split was into two groups roughly corresponding to the 
Autoplusiina + Plusiina and the Euchalciina + Argyrogrammatini. Misidentifications allowed him to 
misplace Macdunnoughia in the Argyrogrammatini, while the other two autoplusiine genera considered, 
Rachiplusia and Autoplusia, were treated together as the closest relatives of Comutiplusia. Chrysanympha 
was mistakenly associated with Plusia, from which it was supposed to have been derived. The Plusiina was 
thus paraphyletic. The topology of the genera of this subtribe was similar to that in Fig. 30 but both Plusia 
and Comutiplusia branched off before Autographa, and Anagrapha was associated with Syngrapha (s.l.). 

Within the Euchalciina (which included Diachrysia), a similar topology to that in Fig. 30 was found. 
However, the branching sequence for the Lamprotes genus-group was inverted and Panchrysia was placed 
with Polychrysia. 

The Argyrogrammatini, with the exception of Anadevidia, was monophyletic (allowing for Macdun- 
noughia, see above), but Chrysodeixis was associated with Argyrogramma rather than with Pseudoplusia. 
Anadevidia was treated as the sister-group of the rest of the Argyrogrammatini -I- Euchalciina. 

Overall, most of the tribes and subtribes (with the exception of the Autoplusiina) are represented 
reasonably accurately on Kostrowicki's phylogenetic scheme. However, their interrelationships and 
several of the generic assignments are at considerable variance with subsequent classifications. 

Ichinose (1962b) (Fig. 2 v Figs 29, 30). Following very shortly after that of Kostrowicki (1961), Ichinose's 
(19626) scheme of Japanese plusiine systematics represented a significant improvement due primarily to 
the inclusion of Abrostola in the study. The recognition of this genus as the most 'primitive' plusiine taxon 
examined resulted in Ichinose reconstructing the first few branch-points within the subfamily more 
accurately than Kostrowicki. Nevertheless, the Argyrogrammatini were paraphyletic, with Chrysodeixis 
(sensu Trichoplusia (s.str.), Chrysodeixis (s.str.) and Ctenoplusia (s.l.)) branching off first, followed by a 
pentachotomy consisting of Ctenoplusia (Acanthoplusia) (as Chrysodeixis) agnata, Thysanoplusia + 
Anadevidia, Diachrysia, the Euchalciina and the Autoplusiina + Plusiina. As for Kostrowicki, this 
interpretation of Ichinose's diagram requires its conversion into cladogram form. The original consisted of 
a series of interconnected circles and thus had numerous extant taxa as putative ancestral forms to other 
extant taxa. Hence some of the branches are zero-length projections (e.g. the branch subtending 
Ctenoplusia (Acanthoplusia) agnata). The Euchalciina and the Plusiina were less well-resolved topologies 
of those found in the present study, while the Autoplusiina (represented by Erythroplusia and Macdun- 
noughia (s.l.) (including Sclerogenia and Antoculeora) was paraphyletic. 

Thus Ichinose's scheme shows a considerable degree of concordance with that proposed here. The 
discrepancies can be largely attributed to his inclusion of only two character complexes in the construction 
of his systematic arrangement - larval chaetotactic 'plan' and the presence/absence of abdominal 
dorso-lateral hair tufts in the males. This accounts for why Chrysodeixis (sensu Ichinose) was biphyletic 
and the Plusiini were not recognized as a monophyletic group (C. ' agnata lacks vestigial larval prolegs on 
segments A3 and A4), and why the Autoplusiina were paraphyletic (both Macdunnoughia [sensu 
Ichinose's 'Scleroplusia'] and Autographa are heterogeneous regarding the presence/absence of seta SV2 
on segment Al). In fact, Ichinose's interconnected circles are probably more correctly interpreted as 
depicting the supposed evolution of plusiine larval chaetotaxy than of the Plusiinae themselves. 

Eichlin & Cunningham (1978) (Fig. 3 v Figs 29, 30). Eichlin & Cunningham's (1978) phylogeny of the 
plusiines is by far the best published approximation to my own results. It incorporated both larval and adult 
characters, with little apparent bias towards either, although the explicit use of some characters was often 
unstated. The result was remarkably congruent with my own. All the tribal and subtribal groups confirmed 
in this study were first recognized by Eichlin & Cunningham, with the exception of the Euchalciina and the 
clade Autoplusiina + Plusiina. 

Thus, the Abrostolini were placed as the most primitive tribe, followed by the basic subdivision of the 
plusias into the Argyrogrammatini and Plusiini . Further analysis of the former was hindered by their virtual 
absence from the Nearctic fauna, only five species from three genera occurring north of Mexico. The 
Autoplusiina were the first to branch off the Plusiini lineage. Rachiplusia and Diachrysia were interpreted 
as the sister-groups of Autoplusia and Allagrapha respectively, conclusions that are consistent with, if not 
entirely supported by, my own results. The branching sequence of Polychrysia, Adeva and Pseudeva was 
identical (allowing for missing Palaearctic taxa) but Chrysanympha and Eosphoropteryx were misplaced as 



SYSTEMATICS OF THE PLUSIINAE 177 

possible branches from the Plusiina. However, Eichlin & Cunningham were unsure of the position of these 
genera. 

The non-monophyletic nature of Autographa, confirmed in the present study (if not in the manner 
originally intended!), was first explicitly noted by Eichlin & Cunningham, although it must be said that this 
conclusion was implicit in Ichinose's (19626) scheme. Eichlin & Cunningham's (1978) arrangement of 
Plusia and Syngrapha (s.l.) agrees more or less with my own, as far as the two are comparable, with the 
exception of their positioning of Anagrapha within Syngrapha. Their conclusions might have been 
different had Macdunnoughia occurred in North America. 

Chou & Lu (1979c) (Fig. 4 v Figs 29, 30). The phylogeny of the Plusiinae proposed by Chou & Lu (1979c) 
represented a retrograde step in the systematics of the Plusiinae . It was overtly phenetic in derivation , very 
poorly resolved and based upon relatively few characters, several of which were of very dubious quality and 
reliability. However, they did recognize Omorphina (Omorphinini) as the most basal plusiine lineage, 
followed by the Abrostolini (Abrostola). If their 'Plusiini' {Diachrysia, Anadevidia, Thysanoplusia) is 
ignored, then the Argyrogrammatini is recovered intact, though it also includes Erythroplasia and 
Puriplusia. The Euchalciina is also regarded as monophyletic. However, the internal relationships of these 
two groups bear only minimal correspondence with my results. Syngrapha (s.l.) was interpreted as a 
lineage independent of the Plusiina + Autoplusiina (less Puriplusia and Erythroplusia) . Thus, despite 
inadequate methods and reliance upon a few tried-and-tested 'classic' characters, several of the most 
robust plusiine clades did emerge intact from Chou & Lu's analysis. 

Kljutshko (1985a, b) (Figs 5-7 v Figs 29, 30). I described in detail above why Kljutshko's original 
cladogram (Fig. 5) of the Palaearctic Plusiinae was erroneous. After correctly applying cladistic methodol- 
ogy, the somewhat more polytomous cladogram (Fig. 6) is comparable with my own findings. Like most 
workers, Kljutshko recovered the Argyrogrammatini (less Anadevidia) as a monophyletic unit. The 
Plusiini were also treated as monophyletic, allowing for the exclusion of Erythroplusia. However, the 
fragmentation of the Euchalciina into four separate lineages resulted in almost complete incongruence 
between Kljutshko's groupings and my own. The only concordant taxa are Syngrapha + Caloplusia and the 
Autoplusiina (represented by Macdunnoughia and Antoculeora). Reanalysis of some of his characters 
(Fig. 7) resulted in the gain of only one more congruent clade, the Argyrogrammatini + Plusiini. Overall, a 
laudable effort at explicit cladistic character analysis but a poor result. 

Discussion 

Larval foodplant associations 

One of the most popular eco-systematic subjects at present is the investigation of the interac- 
tions between herbivorous insects and their foodplants. The plusiines show a marked divergence 
in the number and relationships of the plants that their larvae eat. Of those that are known (a 
proviso that underlies this entire section of the discussion), many plusiine genera are extremely 
polyphagous, often giving the impression that a plant merely has to be green, and preferably 
herbaceous, to be a potential meal. Such genera include Chrysodeixis, Trichoplusia, Thysano- 
plusia, Argyrogramma, Allagrapha, Autoplusia, Anagrapha and Autographa, which include 
most of the notorious agricultural pests of the subfamily. 

At the other extreme, a number of genera are known to be virtually monophagous at the level 
of plant genus. For example, Lophoplusia violacea is known to eat only the labiate Stenogyne, 
while its congener, L. pterylota, is recorded only from Clermontia in the relatively unrelated 
Lobeliaceae. Many other examples were noted above and can be summarized thus: Abrostola 
(Urticaceae and related families, one species on Asclepiadaceae); Mouralia (Commelinaceae); 
Anadevidia (Cucurbitaceae); Ctenoplusia albostriata + putative allies (Asteraceae); Euchal- 
ciina (most) (Ranunculaceae); Euchalcia (part) (Boraginaceae); Chrysanympha (Ericaceae); 
Erythroplusia, Macdunnoughia, Sclerogenia, Antoculeora, Autographa bractea + putative allies 
(all Asteraceae); Plusia (monocotyledons); and Syngrapha celsa and rectangula species-groups 
(conifers) . 

The most striking observation about this wide ranging monophagy and oligiphagy is that there 
is very little pattern to it from the point of view of either plant or plusiine systematics. The 
sister-group of an oligophagous plusiine genus is about as likely to be polyphagous as oligopha- 



178 IAN J. KITCHING 

gous, and if the latter then it will probably feed on a markedly different group of plants. 
Compare, for example, the sister-group pairs of Plusial Lophoplusia, Abrostolal Mouralia and 
Autoplusial Antoculeora . The only two generalizations that can be made are that most euchal- 
ciines feed on Ranunculaceae, and oligophagous autoplusiines seem to show a preference for 
Asteraceae. Many foodplant records are missing for the tropical argyrogrammatines. When 
their life histories are elucidated, I expect equally varied, interesting and oligophagous patterns 
to be found as are presently observed in the remainder of the subfamily. 

Biogeography 

Very little has been published on the biogeography of the Plusiinae, mainly because all of the 
previous studies have examined isolated faunas. This makes a world-wide comparison and 
evaluation virtually impossible, especially from the historical viewpoint. 

Kostrowicki (1961) presented in detail the descriptive and distributional biogeography of the 
plusias, concentrating on the Palaearctic species. He noted that the Argyrogrammatini (his 
group A) were essentially tropical, occurring in the Holarctic region primarily in the southern 
areas. He considered his group B (Autoplusia, Rachiplusia, Cornutiplusia) to be characteristic 
of Notogaea. Notogaea is defined by the Chambers Twentieth Century Dictionary as Australia, 
the islands to the north, New Zealand and Polynesia, while the Concise Oxford English 
Dictionary also includes the neotropics. Neither of these concepts coincides with the range of 
group B, which is absent from Oceania but present also in Africa, and, to a lesser extent, in the 
Holarctic. Kostrowicki's group C (Syngrapha, Caloplusia, Anagrapha) was exclusively Holarc- 
tic (with the exception of one unnamed species in the Andes - presumably Rachiplusia virgula, 
which thus belongs to group B), as was group D (the balance of the plusias), although most 
species of the latter group were Palaearctic. 

Kostrowicki proceeded to examine these distributions in greater detail and speculated on the 
centres of origin and dispersal routes of the various taxa. Group A was supposed to have 
originated in the Old World tropics, while group C was of 'obvious boreal-alpine and superficial- 
ly circum-boreal origin'. Group D was hypothesised to have had its origins in the eastern 
Palaearctic, where most of its species are currently found. However, lack of a consideration of 
the Omorphinini and Abrostolini, coupled with a dispersalist/centre of origin approach resulted 
in much speculative fiction. This was compounded by Kostrowicki's errors of identification and 
his non-critical systematic analysis. 

A better basis for a zoogeographical analysis was Eichlin & Cunningham's (1978) phylogeny. 
However, their comments on the subject comprised merely a single paragraph. Coloured by 
their belief that the Plusiinae were very closely related to the Stictopterinae, they postulated a 
tropical origin for the plusiines, followed by an invasion of the temperate zones. The Argyro- 
grammatini were again recognized as a primarily tropical tribe, the Plusiini as primarily 
temperate, some of which had further adapted to Boreal or Arctic conditions. 

The results of the present study represent the first attempt to analyse the higher classification 
of the Plusiinae on a world-wide basis. It would thus probably be seen as unforgivable if I did not 
make some statements regarding the historical biogeography of the subfamily. However, they 
will necessarily be non-commital and somewhat conjectural, given the ambiguity contained in 
the cladogram upon which they are based. 

In the absence of a reasonable hypothesis as to the immediate outgroup to the Plusiinae, I can 
only speculate where the subfamily might have first arisen. Perhaps the marginally most 
plausible area is the south-eastern Palaearctic/north-eastern Oriental regions. In this area are 
currently found the most plesiomorphic plusiine, Omorphina; the Abrostola species (A 
anophioides) that may represent the intermediate link with Mouralia; several of the more 
plesiomorphic argyrogrammatine genera (notably Anadevidia and Zonoplusia); and some of 
the plesiomorphic members of the three subtribes of the Plusiini: Diachrysia, Erythroplusia, 
Polychrysia. 

If this hypothesis is accepted, for the sake of argument, then the 'proto- Abrostolini' may have 
had a more or less world-wide occurrence, with subsequent vicariance events isolating first 



SYSTEMAT1CS OF THE PLUSIINAE 179 

Mouralia, then the Nearctic abrostolas. A cladistic analysis of the Abrostolini is necessary to test 
this suggestion. 

It appears most likely that the Argyrogrammatini evolved primarily in the tropics and the 
Plusiini in the temperate regions. Superficially, this may suggest a Gondwanan and Laurasian 
distribution pattern respectively. However, it is extremely unlikely that the cladogenetic event 
that ultimately gave rise to the two major plusiine lineages was contemporaneous with the 
primary break-up of Pangea. Such an association would place the time of divergence of the 
Argyrogrammatini and Plusiini at between 180 (Rosen, 1978) and 150 (Irving, 1977) million 
years ago, that is, in the Jurassic, before the currently accepted origin of the angiosperms! A 
more recent origin would seem less incredible but remains unknown. 

There are other interesting patterns of distribution within the subfamily. Chrysodeixis seems 
to show a good example of a transantarctic distribution. C. (Pseudoplusia) is found in South 
America and Saint Helena, while the hypothesised plesiomorphic C. (Chrysodeixis), C. 
argentifera, is found in Australia. More apomorphic Chrysodeixis species are then found 
towards the Equator. This genus would be one of the most profitable to study from a 
biogeographic point of view, as well as containing no fewer than five economically important 
pest species (C. includens, C. argentifera, C. eriosoma, C. chalcites and C. acuta). 

Within the Plusiini, there are several examples of Nearctic/Palaearctic sister-genus pairs: 
Chrysanymphal Panchrysia, Euchalcia (Adeva)l Euchalcia (s.str.), Pseudeval Lamprotes, Alla- 
graphal Macdunnoughia and Autoplusia/Antoculeora. These have only become apparent follow- 
ing a world-wide analysis of the subfamily and all merit further investigation. The conclusion 
that Lophoplusia is the sister-group of Plusia implies that the former, a Hawaiian endemic, 
originated by dispersal from either the eastern Palaearctic or western Nearctic. A decision 
between the two must await the resolution of the interrelationships of the species of these, and 
maybe related, genera. Finally, the most apomorphic syngraphas appear to be those that live 
furthest north, culminating in those with a circumpolar distribution. A cladistic analysis is 
necessary to reveal whether this northward pattern is truly as linear as it first appears. 

All of the above observations are preliminary and tentative, and are meant more as stimuli to 
further research than as statements of fact. More concrete conclusions must await detailed 
cladistic investigations of the individual plusiine genera and a more comprehensive analysis of 
the higher classification. There is no doubt, however, that the Plusiinae could prove to be most 
rewarding subjects for biogeographical analysis. 

Conclusions 

Eichlin & Cunningham (1978) concluded with the hope that their study would be useful as a 
framework for future work. By examining the world fauna, I have endeavoured to synthesize an 
overall classification that has illuminated transcontinental relationships not suspected from the 
previous faunal studies. However, this arrangement is also preliminary, as any first attempt in an 
uncharted field must be . Although I have referred in this paper to over half the plusiine species, I 
have examined in detail less than one-eighth of them. Many species still languish in 'Plusia' s.l. 
and the elucidation of their relationships should have high priority, if only to empty a dustbin. 
Similarly, Trichoplusia ought to be dismembered into its component monophyletic groups but I 
suspect that an analysis of the 'higher' argyrogrammatines may have to be performed first or 
simultaneously, rendering this a massive task. The subtribes of the Plusiini are also poorly 
defined and grey areas often include potential or actual pest species (e.g. Anagrapha falcifera) . 
Autographa, in particular, is in dire need of attention but would have to be analysed in 
conjunction with the rest of the Plusiina. 

There are numerous biological topics that could employ plusiines as subjects. Perhaps the 
most important, as with all other groups of Lepidoptera, is the elucidation of life histories, to 
open up the possibilities for employing early stages in systematic analyses. The plusiines also 
lend themselves to questions of insect/plant interactions (How and why did Abrostola asclepiadis 
switch from Urtica to Vincetoxicuml Why does one Lophoplusia feed on a scentless mint and 
another on a tree lobelia with sticky, milky latex?); of pheromone biology (What does Mouralia 



180 IAN J. KITCHING 

do with its giant hairpencil? Why do Loboplusia and Chrysodeixis have such elaborate scent 
organs?); and of biogeography (Chrysodeixis and Autoplusia are the prime candidates). If 
nothing else, I hope that this study will stimulate interest in the group so that such problems will 
be investigated. The Plusiinae could be much more than gold spots and burnished brasses, 
spectacles and silver Ys. 

Addenda: notes added in proof 

Taxa examined 

Brachionycha sphinx Hufnagel was inadvertently used as the exemplar species for this genus 
instead of the type species, B. nubeculosa Esper. 

Clade analysis 

The larva of Abrostola triopus Hampson has been recorded feeding on Commelina sp. and 
the grass, Pennisetum purpureum (Sevastopulo, 1976). The former record is of particular 
interest as Commelina belongs to the same family (Commelinaceae) as the larval foodplants of 
Mouralia, the sister-genus of Abrostola. A. triopus is found in South Africa (with three doubtful 
records from Ethiopia and Kenya, probably misidentifications) and thus may be a critical species 
in the future understanding of the evolution of the Abrostolini. 

Sevastopulo (1976) also recorded the larva of Trichoplusia vittata (as transfixa) feeding on 
Artemisia africana, so corroborating the hypothesised oligophagy of this group. 

The larval foodplant of Plusiopalpa dichora (as P. adrasta) was given by Sevastopulo (1976) as 
Coleus (Lamiaceae). 

An undescribed species of Euchalcia with yellow hindwings has recently been discovered in 
north-western Canada (J. D. Lafontaine, pers. comm.). 

Polychrysia morigera appears to be extending its range eastwards in the U.S.A. (Covell & 
Medley, 1986). It was recorded from Kentucky in 1976, from Pennsylvania (recent unknown 
date) and was caught for the first time in Tennessee in 1984, trapped in the flowers of a Lady's 
Slipper Orchid (Cypripedium kentuckiense)\ 

Lafontaine (in press) has recently shown that there has been a considerable degree of 
confusion regarding the correct identity of Syngrapha ottolenguii. The type is a female that has 
had the abdomen of a male Autographa calif ornica glued on. Examination of true ottolenguii has 
shown the species to be closely related to S. (Palaeographa) interrogationis and S. (P. ) nyiwonis. 
Thus, S. ottolenguii almost certainly belongs in the subgenus Palaeographa of Syngrapha. In 
addition, the series examined by Eichlin & Cunningham (1978) was apparently mixed. When 
sorted correctly, S. ottolenguii was found to be endemic to Attu Island in the Aleutian Islands. 
'5. ottolenguii 1 in Alaska and Canada was shown by Lafontaine (in press) to be Autographa 
buraetica Staudinger, a species previously known only from the eastern Palaearctic and, until 
recently, considered as a subspecies or even a form of Autographa pulchrina. 

Lafontaine (in press) has also examined a large number of specimens of Syngrapha devergens 
and S. alticola. He concluded that the genitalic differences noted by McDunnough (1944) are 
constant and that the Nearctic S. alticola should be reinstated as a species separate from the 
Palaearctic 5. devergens. 

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Appendix 1 Genera and type species of Plusiinae and outgroups. 

The following is a list of the numbers of each sex examined in the Plusiinae and the outgroups. Shensiplusia 
was scored from drawings published by Chou & Lu (1974), while Desertoplusia and Pseudochalcia were 
described too recently to be included in the main analysis (but see Consensus classification). 

Abrostola Ochsenheimer 1816 
triplasia Linnaeus 1758 2 cf , 2 $ (Fig. 32) 



188 IAN J. KITCHING 

Acanthoplusia Dufay 19706 

tarassota Hampson 1913 2 cf , 2 $ (Fig. 53) 
Adeva McDunnough, 1944 

albavitta Ottolengui 1902 1 Cf , 1 ? (Fig. 71) 
Agrapha Hubner [1821] 1816 

ahenea Hubner [1821] 1816 2 cf , 2 $ (Fig. 57) 
Allagrapha Franclemont 1964 

aerea Hubner [1803] 2 cf , 2 $ (Fig. 75) 
Anadevidia Kostrowicki 1961 

peponis Fabricius 1775 2 cf , 2 $ (Fig. 37) 
Anagrapha McDunnough 1944 

falcifera Kirby 1837 3 cf , 2 $ (Fig. 86) 
Antoculeora Ichinose 1973 

ornatissima Walker 1858 2 cf , 2 $ (Fig. 79) 
Argyrogramma Hubner 1823 

verruca Fabricius 1794 3 cf , 2 § (Fig. 59) 
Autographa Hubner [1821] 1816 

bractea [Denis & Schiffermiiller] 1775 2 cf , 2 $ (Fig. 88) 

gamma Linnaeus 1758 2 cf, 2 $ (Fig. 87) 
Autoplusia McDunnough 1944 

egena Guenee 1852 3 cf , 2 9 (Fig. 80) 
Caloplusia Smith 1884 

hochenwarthi Hochenwarth 1785 2 cf , 2 $ (Fig. 95) 
Chrysanympha Grote 1895 

formosa Grote 1865 1 cf , 1 $ (Fig. 63) 
Chrysodeixis Hubner [1821] 1816 

chalcites Esper 1789 4 cf, 2 $ (Fig. 39) 
Cornutiplusia Kostrowicki 1961 

circumflexa Linnaeus 1767 2 cf , 3 $ (Fig. 89) 
Ctenoplusia Dufay 1970a 

limbirena Guenee 1852 2 cf , 2 $ (Fig. 54) 
Dactyloplusia Chou & Lu 1979 

impulsa Walker 1865 2 cf , 4 $ (Fig. 50) 
Diachrysia Hubner [1821] 1816 

chryson Esper 1789 2 cf , 2 9 (Fig. 84) 
Eosphoropteryx Dyar 1902 

thyatiroides Guenee 1852 2 cf , 2 $ (Fig. 61) 
Erythroplasia Ichinose 1962a 

rutulifrons Walker 1858 4 cf , 2 $ (Fig. 73) 
Euchalcia Hubner [1821] 1816 

variabilis Piller 1783 2 cf , 2 $ (Fig. 67) 
Eutheiaplusia Dufay 1970a 

pratti Kenrick 1917 1 cf (holotype), 1 $ (Fig. 52) 
Lamprotes R.L. 1817 

c-aureum Knoch 1781 2 cf , 2 $ (Fig. 35) 
Loboplusia Roepke 1941 

vanderweelei Roepke 1941 2 cf (holotype (Fig. 74) andparatype), 1 $ (holotype Autographa pokhara) 
Lophoplusia Zimmerman 1958 

pterylota Meyrick 1904 2 cf , 2 $> (Fig. 92) 
Macdunnoughia Kostrowicki 1961 

confusa Stephens 1850 7 cf , 3 $ (Fig. 76) 
Mouralia Walker 1858 

tinctoides Guenee 1852 2 cf , 2 $ (Fig. 33) 
Omorphina Alpheraky 1892 

aurantiaca Alpheraky 18921 cf, 1 $ (Fig. 31) 
Palaeographa Kljutshko 1984 

interrogationis Linnaeus 1758 2 cf , 2 $ (Fig. 93) 
Panchrysia Hubner [1821] 1816 

deaurata Esper 1787 2 cf , 2 $ (Fig. 64) 



SYSTEMATICS OF THE PLUSIINAE 189 



Plusia Ochsenheimer 1816 

festucae Linnaeus 1758 2 cf , 2 9 (Fig. 90) 
Plusidia Butler 1879 

c/i«'ra/itfiiTauscher 1809 2 d", 2 $ (Fig. 34) 
Plusiopalpa Holland 1894 

dichora Holland 1894 1 cf , 2 $ (Fig. 58) 
Plusiotricha Holland 1894 

livida Holland 1894 1 cf 
Polychrysia Hiibner [1821] 1816 

moneta Fabricius 1787 2 cf , 2 $ (Fig. 62) 
Pseudeva Hampson 1913 

purpurigera Walker 1858 2 cf , 2 9 (Fig. 36) 
Pseudoplusia McDunnough 1944 

includens Walker 1857 4 cf , 2 $ (Fig. 38) 
Puriplusia Chou & Lu 1974 

purissima Butler 1878 2 cf , 2 ? (Fig. 77) 
Rachiplusia Hampson 1913 

nu Guenee 1852 4 cf , 2 9 (Fig. 72) 
Sclerogenia Ichinose 1973 

Jessica Butler 1878 3 cf , 2 $ (Fig. 78) 
Shensiplusia Chou & Lu 1974 

nigribursa Chou & Lu 1974 [1 9] 
Stigmoplusia Dufay 1970a 

chalcoides Dufay 1968 1 Cf , 1 9 (Fig. 60) 
Syngrapha Hiibner [1821] 1816 

devergens Hiibner [1813] 2 cf , 2 9 (Fig. 94) 
Thysanoplusia Ichinose 1973 

intermixta Warren 1913 3 cf , 2 9 (Fig. 47) 
Trichoplusia McDunnough 1944 

ni Hiibner [1803] 3 cf , 2 9 (Fig. 43) 
Zonoplusia Chou & Lu 1979 

ochreata Walker 1865 3 cf , 2 9 (Fig. 49) 

Plusiine taxa described too recently for inclusion in the analysis 

Desertoplusia Kljutshko 1985 

bella Christoph 1887 (Fig. 70) 
Pseudochalcia Kljutshko 1985 

inconspicua Graeser 1892 (Fig. 69) 

Outgroups (nominal subfamily given in parentheses) 

Anuga Guenee, 1852 

constricta Guenee 1852 (Euteliinae) 2 cf , 2 9 (Fig. 101) 
Brachionycha Hiibner [1819] 1816 

sphinx Hufnagel 1766 (Cuculliinae) 2 cf , 2 9 (Fig. 106) 
Calophasia Stephens 1829 [June] 

lunula Hufnagel 1766 (Cuculliinae) 2 cf , 2 9 (Fig. 104) 
Chalcopasta Hampson 1908 

territans Edwards 1884 (Amphipyrinae) 2 cf , 2 9 (Fig. 102) 
Cucullia Schrank 1802 

umbratica Linnaeus 1758 (Cuculliinae) 2 cf , 2 9 (Fig. 105) 
Diloba Boisduval 1840 

caeruleocephala Linnaeus 1758 (Dilobidae) 2 cf , 2 9 (Fig. 107) 
Lophoptera Guenee 1852 

squammigera Guenee 1852 (Stictopterinae) 2 cf , 3 9 (Fig. 99) 
Magusa Walker 1857 

orbifera Walker 1857 (Amphipyrinae) 2 cf , 2 9 (Fig. 97) 
Oncocnemis Lederer 1853 

confusa Freyer [1840] (Cuculliinae) 2 cf , 2 9 (Fig. 103) 



190 IAN J. KITCHING 

Paectes Hiibner 1818 

cristatrix Guenee 1852 (Euteliinae) 3 cf , 2 $ (Fig. 100) 
Stictoptera Guenee 1852 

cucullioides Guenee 1852 (Stictopterinae) 2 cf , 4 9 (Fig. 98) 

Appendix 2 Plusiine generic-level taxa not employed in this study. 

Cerviplusia Chou & Lu 1974. Type species: Cerviplusia wukongensis Chou & Lu 1974. C. wukongensis was 
synonymized with Antoculeora ornatissima (as Macdunnoughia ornatissima) by Dufay (1977). Thus, 
Cerviplusia is a junior subjective synonym of Antoculeora Ichinose. 

Chrysaspidia Hiibner [1821] 1816. A junior objective synonym of Plusia Ochsenheimer. 

Chrysoptera Berthold 1827. A junior objective synonym of Lamprotes R.L. 

Cubena Walker 1856. Type species: Phalaena polydamia Stoll 1782. C. polydamia is a junior subjective 
synonym of Lamprotes c-aureum and has been treated as such for about a century. Cubena is thus a 
junior subjective synonym of Lamprotes. 

Inguridia Butler 1879. Type species: Inguridia abrostolina Butler 1879. A junior subjective synonym of 
Abrostola Ochsenheimer. There is no doubt that abrostolina is congeneric with Abrostola triplasia and as 
far as I am aware, Inguridia has not been used this century. 

Melanoplusia Chou & Lu 1978. A junior objective synonym of Sclerogenia Ichinose. 

Neoplusia Okano 1963. Type species: Neoplusia furahatai Okano 1963. N. furahatai was synonymized with 
Chrysodeixis acuta Walker by Dufay (1965), based on an examination of the genitalia. Neoplusia is thus a 
junior subjective synonym of Chrysodeixis Hiibner and has been treated as such since 1965. 

Palaeoplusia Hampson 1913. Type species: Plusia venusta Walker 1865. McDunnough (1944) considered 
P. venusta to be congeneric with Plusia festucae (as Chrysaspidia festucae). He therefore synonymized 
Palaeoplusia with Chrysaspidia and thus the former became a junior subjective synonym of Plusia 
Ochsenheimer. This decision has been upheld since and is followed in this study. 

Paraplusia Mukerji & Krishnamorthy 1955. Type species: Plusia confusa Stephens 1850. This name is not 
nomenclaturally available because it was published after 1930 without a statement of characters 
differentiating the taxon. 

Perloplusia Chou & Lu 1978. Type species: Perloplusia pseudopyropia Chou & Lu 1978. Chou & Lu 
(1979a) subsequently considered this species to be synonymous with Erythroplusia pyropia Butler and 
thus, by implication, Perloplusia is a junior subjective synonym of Erythroplusia (syn. n.). 

Podioplusia Ichinose 1962a. A junior objective synonym of Anadevidia Kostrowicki. 

Scleroplusia Ichinose 1962a. A junior objective synonym of Macdunnoughia Kostrowicki. 

Unca Oken 1815. Included in a work rejected for nomenclatural purposes (ICZN, 1956). 

Unca Lhomme [1929]. A junior objective synonym of Abrostola Ochsenheimer. 

Yerongponga Lucas 1901. Type species: Yerongponga exequialis Lucas 1901. This genus and species was 
described in the Plusidae [sic] at a time when Plusidae referred to the quadrifine noctuids in general 
(Kitching, 1984). Yerongponga then, by inertia, became included in the Plusiinae (Nye, 1975). The 
species was not examined in this study (the type specimen could not be located; it was searched for, but 
not found, in the National Insect Collection, CSIRO, Canberra and in the South Australia Museum, 
Adelaide). However, from the description, the species is evidently much larger than any known plusiine 
and is most likely to be a species of Platyja Hiibner (J. D. Holloway, pers. comm.). Thus Yerongponga 
can be transferred to the Catocalinae. 









SYSTEMATICS OF THE PLUSIINAE 

Appendix 3 Data matrix 



191 



Data set, characters 1-70, Plusiinae. 

Omorphina 00000 01000 00101 00100 01010 OHIO 10000 00000 01000 0***0 01000 10010 00000 01000 



Abrostola 


11000 


00001 


00101 


00101 


02000 


01010 


10000 


00000 


00000 


0**10 


10000 


10100 


00000 


01000 


Moural ia 


11000 


00001 


00122 


00211 


01000 


01010 


10000 


00000 


00000 


0**10 


10000 


10100 


00030 


01000 


Anadevidia 


00010 


00001 


00111 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00000 


01000 


Dactyloplusia 


00110 


10001 


01121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10111 


00000 


01000 


Pseudoplusia 


00110 


10001 


11121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00000 


01000 


Chrysodeixis 


00110 


10001 


10121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00000 


01000 


Zonoplusia 


00100 


10001 


01121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10000 


00000 


01000 


Trichoplusia 


00110 


10001 


01121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10101 


00000 


01000 


Thysanoplusia 


00010 


10011 


01121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10101 


00000 


01000 


Plusiotricha 


00110 


10001 


101*1 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10102 


10000 


01000 


Eutheiaplusia 


00100 


10001 


00121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00000 


01000 


Agrapha 


00110 


00001 


01121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00000 


01000 


Acanthoplusia 


00120 


10001 


00121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10101 


00000 


01000 


Ctenoplusia 


00130 


10001 


00121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10102 


00000 


01000 


Plusiopalpa 


00110 


10001 


01121 


00001 


01000 


01010 


10000 


00000 


10000 


0**10 


00000 


10100 


10000 


01000 


Argyrogramma 


00110 


10001 


00121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10103 


30000 


01000 


Stigmoplusia 


00110 


10001 


10121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10103 


20000 


01000 


Shensiplusia 


001** 


*0001 


*11*1 


*0**1 


01000 


01010 


10000 


000*0 


00000 


0**10 


00000 


10*** 


***** 


01000 


Eosphoropteryx 


00110 


10001 


00101 


00101 


01000 


01010 


10000 


00010 


00000 


0**10 


00000 


10*10 


00000 


01000 


Polychrysia 


00111 


10001 


00101 


00101 


01000 


01010 


10000 


00010 


10000 


0**11 


00000 


10110 


00002 


01000 


Panchrysia 


00010 


10001 


00101 


10111 


01000 


01010 


10000 


00010 


01000 


0**10 


00000 


10110 


00003 


01000 


Chrysanympha 


00011 


10001 


00101 


00101 


01000 


01010 


10000 


00010 


11000 


0**10 


00000 


10*10 


00000 


01000 


Euchalcia 


00040 


00001 


00101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10010 


00000 


01000 


Adeva 


00041 


00001 


00101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10*00 


00000 


01000 


Plusidia 


00010 


00001 


00101 


00101 


01000 


01010 


10000 


00010 


00000 


0**10 


00000 


10010 


00000 


01000 


Lamprotes 


00110 


10001 


00101 


10111 


01000 


01010 


10000 


00010 


11000 


0**10 


00000 


10110 


00000 


01000 


Pseudeva 


00011 


10001 


00101 


10111 


01000 


01010 


10000 


00010 


11000 


0**11 


00000 


10110 


00000 


01000 


Rachiplusia 


00110 


10001 


00121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00120 


01000 


Erythroplusia 


00111 


00001 


00101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00000 


01000 


Al lagrapha 


00010 


10001 


00101 


10101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10010 


00000 


01000 


Macdunnoughia 


00110 


10001 


00101 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00000 


01000 


Puriplusia 


00110 


10001 


00101 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00000 


01000 


Loboplusia 


00110 


00001 


01101 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00000 


01000 


Sclerogenia 


00110 


10001 


01101 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00000 


01000 


Autoplusia 


00110 


10001 


00121 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00010 


01000 


Antoculeora 


00110 


10001 


01101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00000 


01000 


Diachrysia 


00010 


10011 


00101 


10101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10010 


00001 


01000 


Anagrapha 


00110 


10001 


00101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00020 


01000 


Autographa gamma 


00110 


10001 


01101 


00001 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00000 


01000 


A. bractea 


00110 


10001 


00101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00000 


01000 


Cornutiplusia 


00110 


00001 


01101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00010 


01000 


Plusia 


00110 


00001 


00101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10010 


00000 


01000 


Lophoplusia 


00110 


00001 


01101 


00001 


01000 


01010 


10000 


00000 


00000 


21110 


00000 


10100 


00003 


01000 


Palaeographa 


00110 


10001 


01101 


00101 


01000 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00010 


01000 


Syngrapha 


00110 


01001 


01101 


00101 


01010 


01010 


10000 


00000 


00000 


0**10 


00000 


10100 


00010 


01000 


Caloplusia 


00110 


01001 


01101 


00101 


01010 


01010 


10000 


00000 


00000 


0**10 


00000 


10110 


00220 


01000 



192 IAN J. KITCHING 

Data set, characters 71-150, Plusiinae. 

Omorphin 00000 00**0 ***** ***** ***** 00000 00000 00000 00000 00100 00000 01001 10000 01000 00310 *1100 

Abrostol 00100 00**1 **0** ***0* 00000 00011 01100 01000 00000 001*0 2*011 1000* 0*010 00000 001** 00100 

Mouralia 00100 00**1 **500 00000 00000 00011 01100 01000 00000 00100 00011 1000* 0*010 10000 0020* 00100 

Anadev 00200 00**1 **300 00010 00000 00010 00001 10100 00000 10100 00000 00001 10000 10000 00310 000** 

Oactylo 00200 11102 **200 00010 11120 00010 00001 10000 00000 00100 00000 00101 10000 10000 00310 00110 

Pseudo 00200 10**1 **0** ****0 00000 00010 00001 10001 10000 00101 00000 00200 10000 10000 00300 00100 

Chryso 00200 13001 **500 00010 10110 01010 00000 10001 10000 00101 00000 01100 10000 10000 00300 00100 

Zono 00200 10**1 **200 00010 10020 00010 00000 10101 00000 00100 00000 00001 10000 10000 00310 00100 

Tricho 00200 11002 **310 00010 11220 00010 00001 10100 00000 00100 00000 0000* 0*000 10000 00300 00110 

Thysano 00200 11002 **300 00010 11120 00010 00000 10101 01000 00100 00000 00101 10000 10000 00300 00100 

P'tricha 00200 12002 **210 00010 21220 00010 00000 10001 03000 00110 00000 0*001 10100 10101 00320 00100 

Eutheia 00200 12002 **0** ***10 ****0 00010 00000 10001 00000 00100 00000 0*001 10000 10101 00310 00100 

Agrapha 00200 12012 **210 11110 21220 00110 00001 10101 01000 00100 00000 00000 10000 10100 00320 00110 

Acantho 00200 11002 **300 11010 23220 00010 00000 10001 01000 00110 00000 00001 10100 10100 00300 00100 

Cteno 00200 11202 **210 01010 23220 00010 00000 10101 01000 00110 00000 00001 10100 10100 00300 00100 

P'palpa 00200 12002 **400 00010 13220 00010 00000 10101 01000 00100 00000 00101 10000 10000 00300 00100 

Argyro 00200 12332 **411 10111 22330 00110 00001 10001 01000 00100 20000 00000 10000 10000 00300 00100 

Stigmo 00200 12022 **311 10011 22330 01010 00000 10101 12000 00100 00000 00000 10000 10*00 00310 00100 

Shensi 00200 ***** ***** ***** ***** ***** ***** ***** ***** ***** ***** ***** ***** ***** ***** ***** 

Eosphor 00200 00**1 **0** ***0* 00000 00010 10000 10000 00000 01100 10000 01001 10000 00000 00300 00100 

Polychry 00200 00**1 **0** ***0* 00000 00010 10000 10010 00000 01100 10000 01001 10000 00000 00300 *1100 

Panchry 00200 00**1 **0** ***0* 00000 00010 10000 10010 00000 01100 10000 01001 11000 00000 00300 *1100 

Chrysan 00200 00**1 **0** ***0* 00000 00010 10000 11010 00000 01100 10000 01001 11000 00000 00300 *1100 

Euchalc 00200 00**1 **0** ***0* 00000 00010 10000 10010 00000 01100 10000 01001 10000 00000 00310 *1100 

Adeva 00200 00**1 **0** ***0* 00000 00010 10000 10010 00000 01100 10000 01001 10000 00000 00310 *1100 

Plusid 00200 00**1 **0** ***0* 00000 00010 10000 10010 00000 01100 10000 01001 10000 00000 00320 *1100 

Lamprot 00200 00**1 **0** ***0* 00000 00010 10000 10010 00000 01100 10000 01001 10000 00000 00310 *1100 

Pseudev 00200 00**1 **0** ***0* 00000 00010 10000 10010 00000 01100 10000 01001 10000 00000 00320 *1100 

Rachi 00200 00**1 **300 00000 00000 00010 00000 10001 00010 11100 10000 01001 10000 00010 00300 000** 

Erythro 00200 00**1 **200 00000 00001 00010 00010 10100 00000 11100 10000 01001 10000 00000 10301 001** 

Alia 00200 00**1 **200 00000 00001 00020 00000 10000 00100 11100 10000 00001 10000 01010 11300 00100 

Macdunn 00200 00**1 **200 00000 00001 00020 00000 10000 00100 11100 10000 01001 10000 01010 11301 00102 

Puri 00200 00**1 **200 00000 00001 00020 00000 10000 00100 11100 10000 01001 10000 01010 11301 00101 

Lobo 00200 0***1 **200 0**00 00001 00010 *000* 10000 00100 *1100 100** 0**01 10000 000*0 00310 000** 

Sclero 00200 00**1 **200 00000 00000 00010 00000 10100 00000 11100 10000 01001 10000 00000 12300 000** 

Autoplus 00200 00**1 **300 00000 00001 00010 00000 10000 00000 11100 10000 0100* 0*000 00000 02300 000** 

Antocul 00200 00**1 **200 00000 00001 00010 00010 10000 00000 00200 10000 01001 10000 00*0* 12301 000** 

Diachry 00200 00**1 **100 00000 00000 00010 00000 10000 00000 11100 10000 01001 10000 00000 00300 1*100 

Anagraph 00200 00**1 **100 00000 00000 00010 00001 10000 00100 11100 10000 02001 10000 00000 00210 1*100 

A. gamma 00200 00**1 **100 00000 00000 00010 00001 10110 00000 11100 10000 01001 10000 00000 00300 1*100 

A. bract 00200 00**1 **100 00000 00000 00010 00001 10110 00000 11100 10000 01001 10000 00000 00300 1*100 

Cornuti 00200 00**1 **100 00000 00000 00010 00001 10110 00000 11100 10000 01001 10000 00000 00300 1*100 

Plusia 00200 00**1 **300 00000 00000 00010 00000 10110 00000 11100 10000 01001 10000 00000 00300 1*100 

Lopho 00200 00**1 **100 00000 00000 00010 00001 11110 00000 11100 10000 01001 10000 00000 00320 1*100 

Palaeo 00200 00**1 **100 00000 00000 01010 00001 10110 00000 11100 10000 02001 10000 00000 00310 1*110 

Syngraph 00200 00**1 **0** ***0* 00000 01010 00001 10110 00000 11100 10000 03001 10000 00000 00310 1*110 

Caloplus 00200 00**1 **100 00000 00000 01010 00001 10110 00000 11100 10000 03001 10000 00000 00310 1*110 



SYSTEMATICS OF THE PLUSIINAE 193 

Data set, characters 151-216, Plusiinae. 

Omorphin 00000 01000 00*** *0000 00000 00100 01010 00000 00000 00000 00000 00000 00200 

Abrostol 00000 01000 00*** *0021 00000 00100 01000 01100 00000 00000 00000 00000 00000 
Mouralia 00000 01000 00*** *0011 00000 00100 01000 01100 10000 00000 00000 00000 00000 

Anadev 00000 02200 01000 00000 00**0 00100 01010 00000 00000 00000 00000 00010 00000 

Dactyl o 00000 02200 00*** *1000 01110 00100 01011 00000 00000 00000 00000 00010 00000 

Pseudo 00000 02200 00*** *1000 OHIO 00100 01011 00000 00000 10000 00000 00000 00010 

Chryso 00000 02200 00*** *1000 00110 00100 01011 00000 00000 10000 00000 01010 00000 

Zono 00000 02200 01000 01000 00100 00100 01011 00000 00000 00000 00110 00010 00000 

Tricho 00000 02200 01000 01000 00100 00100 01011 00000 00000 00000 00211 00000 00000 

Thysano 00000 02210 01000 01000 00100 00100 01011 00000 00000 00000 00110 00000 00000 

P'tricha 00000 02211 00*** *1000 00100 **l** ***** ***** ***** ***** ***** ***** ***** * 

Eutheia 00000 02211 00*** *1000 00100 00100 01011 00000 00000 00000 00111 00010 00000 

Agrapha 00000 02200 01000 01000 00100 00100 01011 00000 00000 10000 00000 01000 00010 

Acantho 00000 02200 00*** *1000 00100 00100 01011 00000 00000 01000 00000 00000 00000 

Cteno 00000 02200 00*** *1000 00100 00100 01011 00000 00000 01000 00000 00000 00010 

P'palpa 00000 02200 00*** *1000 00100 00100 01011 00000 00000 00000 10000 00000 00210 

Argyro 00000 02200 01000 01000 00100 00100 01011 00000 00000 00000 00000 00000 00010 

Stigmo 00000 02200 01000 01000 00100 00100 01011 00000 00000 00000 00000 00000 00000 

Shensi ***** ***** ***** ***** ***** ***00 010** *0000 0*000 10*00 001*0 000*0 000*0 * 

Eosphor 00000 02100 00*** *0000 00001 00200 01010 00000 00000 00000 00000 00000 00100 

Polychry 00000 02100 01001 00000 00000 00200 01010 00000 00000 10000 00000 00000 00000 

Panchry 00010 02100 01100 00000 00**0 00200 OHIO 00000 00000 10000 00000 00010 00000 

Chrysan 00010 02100 01100 00000 00**0 00200 OHIO 00000 00000 10000 00000 00000 00000 

Euchalc 00010 02100 01010 00100 00000 00200 01010 00000 00000 10000 10001 00000 00000 

Adeva 00010 02100 01010 00100 00000 00200 OHIO 00000 00000 10000 00001 00000 00000 

Plusid 00000 02100 01000 00100 10**0 00200 OHIO 10000 00020 10000 00001 00000 00000 

Lamprot 00000 02100 00*** *0100 10**0 00200 OHIO 10000 10010 10000 10000 00000 00000 

Pseudev 00000 02100 01000 00100 10**0 00200 OHIO 10000 00020 10000 00000 00000 00000 

Rachi 00010 02100 01000 00000 00000 00100 01010 00000 10000 00300 00000 10000 00200 1 

Erythro 00001 02200 00*** *0000 00**0 00100 01010 00010 00100 00000 00000 00010 00000 

Alia 00000 02100 00*** *0000 00000 11100 01010 00010 10100 00000 10000 00000 00200 

Macdunn 00000 02200 00*** *0000 00**1 11100 01010 00010 11100 00000 00000 10000 00300 

Puri 00000 02100 00*** *0000 00000 10100 01010 00010 12100 00000 00000 10010 00300 

Lobo 0000* 02100 00*** *0*0* 00**1 01*00 010*0 00010 *0100 00000 00000 00010 00200 

Sclero 00001 02100 00*** *0000 00000 11100 01010 00010 00100 00100 00000 00000 01300 

Autoplus 00000 02000 00*** *0000 00000 11100 01010 00010 00100 00211 00000 00000 01300 1 

Antocul 00000 02000 00*** *0000 00000 11100 01010 00000 10100 00211 00000 00000 01300 

Diachry 00000 02100 01001 00000 00000 00100 01010 00000 10000 00000 00000 00000 00000 

Anagraph 00000 02100 01000 00000 00000 00100 01010 00010 00000 00000 00000 00000 00000 

A. gamma 00000 02100 01000 00000 00000 00100 01010 00000 00000 00000 00000 00000 00000 

A. bract 00000 02100 01000 00000 00**0 00100 01010 00000 00000 00000 00000 00000 00002 

Cornuti 00000 02100 01000 00000 00**0 00100 01010 00000 00000 00000 00000 00000 00003 

Plusia 00000 02100 01000 10000 00000 00100 01010 00000 00000 00000 00000 00000 00001 

Lopho 00000 02100 01000 20000 00000 00100 01010 00000 00000 00000 00000 00000 00001 

Palaeo 00000 02100 00*** *0000 00000 00100 01010 00010 10000 00000 10000 00000 00000 

Syngraph 00000 02100 00*** *0000 00000 00100 01010 00010 10000 00000 10000 00000 00000 

Caloplus 00000 02100 00*** *0000 00**0 00100 01010 00010 10000 00000 10000 00000 00000 



194 



IAN J. KITCHING 



Data set, characters 1-70, 71-140, 141-210, 211-216, outgroups. 

Magusa 00000 00000 00102 00312 00100 10100 21111 01000 00000 0**00 00000 10*00 00000 01000 

Stictoptera 10000 00100 00050 01002 00101 10101 21111 01000 00101 0**00 00100 11*00 12000 11101 

Lophoptera 00000 00100 00050 01002 00101 10100 21111 01000 00100 0**00 02000 11000 12000 11001 

Paectes 00000 00000 00040 00002 00101 10100 21212 11000 00101 10100 02000 11100 01000 11201 

Anuga 10000 00000 00030 01002 00101 10101 21212 11000 00000 10100 02100 11100 01000 11201 

Chalcopasta 00000 00011 00002 00201 00000 00100 21101 00001 00010 0**10 00000 00110 00000 01010 

Oncocnemis 00000 00000 00102 00201 01000 00100 21101 00001 01010 0**00 00000 00000 00000 00010 

Calophasia 00000 00000 00103 00311 01000 01100 20101 00100 00000 0**00 00010 00000 00000 01000 

Cucullia 00000 00000 00102 00111 02000 00100 20001 00101 00000 0**00 00010 00*10 00000 01000 

Brachionycha 00000 00000 00122 00100 10000 00100 02*2* 00101 00010 22000 00001 00000 00000 01010 

Diloba 00000 00000 00161 00000 12000 02100 02*2* 00101 00000 22000 00001 00*00 00000 00000 



Magusa 

Stictoptera 

Lophoptera 

Paectes 

Anuga 

Chalcopasta 

Oncocnemis 

Calophasia 

Cucullia 

Brachionycha 

Diloba 



00001 00**3 
01300 00**3 
00300 00**3 
11310 00**3 
11310 00**3 
00000 00**0 
00000 00**0 
00000 00**3 

00002 00**3 
10000 00**0 
10000 00**0 



***** 
***** 
***** 
***** 
***** 
***** 
***** 



***** 
***** 
***** 
***** 
***** 
***** 
***** 



It*** ***** 



00000 
00000 
00000 
1***0 
2***0 
01000 
00000 
1***0 
00000 
1***0 
1***0 



00000 
00000 
00000 
0**00 
0**00 
00001 
00000 
0**00 
00000 
0**00 
0**00 



0100* 
0100* 
01000 
*0001 
*0000 
10000 
00000 
00000 
00010 
00000 
00000 



00000 
000*0 
00010 
00011 
00011 
00000 
00000 
00000 
00000 
00000 
00000 



00000 
0***0 
0***0 
00*00 
0**00 
00100 
001*0 
000*0 
00100 
00100 
0***0 



00000 
10100 
10100 
01100 
01100 
00000 
3*000 
3*000 
00000 
00000 
**100 



0000* 
000** 
0000* 
0001* 
0001* 
0000* 
0000* 
0000* 
0010* 
0000* 
0000* 



0*001 
0*000 
0*000 
0*000 
0*000 
0*001 
0*001 
0*001 
0*001 
0*000 
0*000 



00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 



Magusa 

Stictoptera 

Lophoptera 

Paectes 

Anuga 

Chalcopasta 

Oncocnemis 

Calophasia 

Cucul 1 ia 

Brachionycha 

Diloba 



010** 
000** 
010** 
000** 
000** 
001** 
001** 
010** 
0020* 
000** 
100** 



00100 
000** 
00100 
000** 
000** 
00100 
00100 
00100 
00100 
00100 
00100 



00000 
10100 
10100 

on** 
on** 

00000 
00000 
00000 
00000 
00000 
00000 



10000 00*** 
11000 10*** 
11000 10*** 
00000 10*** 
10000 10*** 
01000 00*** 
01000 00*** 
01000 00*** 
01000 00*** 
01000 00*** 
01000 00*** 



*0000 
**000 

**ooo 

*0000 

**ooo 

*0000 
*0000 
*0000 
*0000 
*0000 
*0000 



00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 



00000 
00111 
00111 
00111 
00111 
00000 
00000 
00001 
00000 
10000 
00000 



00000 
10*00 
10*00 
00*00 
00*00 
00000 
00000 
00000 
00000 
00000 
00000 



00000 
00000 
00001 
00001 
00001 
00000 
00000 
00000 
00000 
00000 
00000 



00000 
00000 
00001 
00001 
00001 
00000 
00000 
00000 
00000 
10000 
00020 



00000 
01000 
000*0 
000*0 
000*0 
10000 
00000 
00000 
10000 
00000 
10000 



00000 
00000 
01000 
01000 
01000 
00000 
00000 
00000 
00001 
00000 
00000 



00001 
00002 
00022 
00002 
00002 
00000 
00010 
00001 
00011 
00100 
00100 



Magusa 

Stictoptera 

Lophoptera 

Paectes 

Anuga 

Chalcopasta 

Oncocnemi s 

Calophasia 

Cucullia 

Brachionycha 

Diloba 



00000 
10000 
20000 
10000 
00000 
00200 
00000 
00003 
00000 
00000 
00200 



SYSTEMATICS OF THE PLUSIINAE 195 

Appendix 4 New taxa and synonymy, status changes and revivals. 

New taxa 

Autoplusiina subtrib. n. 
New combinations (previous generic combination given in parentheses, genus of original description given 
in square brackets). 

Pseudoplusia dalei (Wollaston) (from 'Plusia' dalei) [Plusia] 

Agrapha meretricia (Schaus) (from 'Plusia' meretricia) [Plusia] 

Agrapha calceolaris (Walker) (from 'Plusia' calceolaris) [Plusia] 

Autographa camptosema (Hampson) (from Macdunnoughia camptosema) [Phytometra] 

Autographa schalisema (Hampson) (from Macdunnoughia schalisema) [Phytometra] 
New synonymy 

Autographa emmetra Dufay, syn. n. of A. schalisema (Hampson) 

Perloplusia Chou & Lu, syn. n. of Erythroplusia Ichinose 
Valid genera 

Thysanoplusia gen. rev. (from synonymy with Trichoplusia) 

Ctenoplusia gen. rev. (from synonymy with Agrapha) 

Cornutiplusia gen. rev. (from synonymy with Syngrapha) 

Lophoplusia stat. n. (from subgenus of Plusia s.l.) 
Subgeneric changes 

Pseudoplusia stat. n. - subgenus of Chrysodeixis 

Eutheiaplusia stat. n. - subgenus of Plusiotricha 

Acanthoplusia stat. rev. - subgenus of Ctenoplusia 

Adeva stat. n. - subgenus of Euchalcia 

Caloplusia stat. n. - subgenus of Syngrapha 



Appendix 5 Generic classification of the Plusiinae 

Subfamily Plusiinae 
Tribe Omorphinini 

Omorphina 
Tribe Abrostolini 
Abrostola 
Mouralia 
Tribe Argyrogrammatini 

Argyrogrammatini incertae sedis: Shensiplusia 
Anadevidia 
Chrysodeixis 
subgenus Chrysodeixis 
subgenus Pseudoplusia stat. n. 
Zonoplusia 

Argyrogramma-complex 
Trichoplusia, sedis mutabilis 
Thysanoplusia genus-group, sedis mutabilis 
Thysanoplusia stat. rev. 
Dactyloplusia 
Plusiotricha, sedis mutabilis 
subgenus Plusiotricha 
subgenus Eutheiaplusia stat. n. 
Ctenoplusia, sedis mutabilis stat. rev. 
subgenus Ctenoplusia stat. rev. 
subgenus Acanthoplusia stat. rev. 
Agrapha, sedis mutabilis 
Plusiopalpa , sedis mutabilis 
Argyrogramma genus-group, sedis mutabilis 
Argyrogramma 
Stigmoplusia 



196 IAN J. KITCHING 

Tribe Plusiini 

Subtribe Euchalciina 

Euchalciina incertae sedis: Pseudochalcia 
Eosphoropteryx 
Polychrysia 

Panchrysia genus-group, sedis mutabilis 
Panchrysia 
Chrysanympha 
Euchalcia genus-group, sedis mutabilis 
Desertoplusia 
Euchalcia 

subgenus Euchalcia 
subgenus Adeva 
Lamprotes genus-group, sedis mutabilis 
Plusidia 
Lamprotes 
Pseudeva 
Plusiini incertae sedis: Diachrysia 
Anagrapha 
Rachiplusia 
Subtribe Autoplusiina subtrib. n. 

Autoplusiina incertae sedis: Erythroplusia 

Loboplusia 
Autoplusia genus-group 
Sclerogenia 
Autoplusia 
Antoculeora 
Macdunnoughia genus-group 
Allagrapha 
Macdunnoughia 

subgenus Macdunnoughia 
subgenus Puriplusia 
Subtribe Plusiina 
Autographa, sedis mutabilis 
Plusia-comp\ex, sedis mutabilis 
Plusia genus-group 
Plusia 

Lophoplusia stat. n. 
Cornutiplusia genus-group 
Cornutiplusia stat. rev. 
"Autographa" bractea 
Syngrapha, sedis mutabilis 
subgenus Palaeographa 
subgenus Syngrapha 
subgenus Caloplusia stat. n. 



SYSTEMATICS OF THE PLUSIINAE 



197 



31 







* 



34 





. t._v 




35 



36 



Figs 31-36 31, Omorphina aurantiaca (wingspan [ws] = 22 mm). 32, Abrostola triplasia (ws = 33 mm). 
33, Mouralia tinctoides (ws = 45 mm). 34, Plusidia cheiranthi (ws = 30 mm). 35, Lamprotes c-aureum (ws 
= 33 mm). 36, Pseudeva purpurigera (ws = 32 mm). Wingspans for Figs 31-107 are measured from 
wingtip to wingtip and are given to the nearest mm. 






198 



IAN J. KITCHING 





38 





40 




41 




Figs 37-42 37, Anadevidia peponis (ws = 41 mm). 38, Chrysodeixis (Pseudoplusia) includens (ws = 37 
mm). 39, Chrysodeixis (Chrysodeixis) chalcites (ws = 38 mm). 40, C. (C. ) argentifera (ws = 35 mm). 41, 
C. (C.) illuminata (ws = 39 mm). 42, C. (C) kebea (ws = 34 mm). 



SYSTEMATICS OF THE PLUSIINAE 



199 





\ 










I^S 




& 








43 












47 




%jm 






■ 



48 



Figs 43-48 43, Trichoplusia ni (ws = 35 mm). 44, T. sestertia (ws = 30 mm). 45, T. exquisita (ws = 35 
mm). 46, T. lectula (ws = 30 mm). 47, Thysanoplusia intermixta (ws = 35 mm). 48, Thy. daubei (ws = 27 
mm). 



200 



IAN J. KITCHING 







51 



52 




53 



f 




Figs. 49-54 49, Zonoplusia ochreata (ws = 25 mm). 50, Dactyloplusia impulsa (ws = 24 mm). 51, 
Plusiotricha (Plusiotricha) sp. near ftWda (ws = 31 mm). 52, Plusiotricha (Eutheiaplusia) pratti (ws = 36 
mm). 53, Ctenoplusia (Acanthoplusia) tarassota (ws = 35 mm). 54, Ctenoplusia (Ctenoplusia) limbirena 
(ws = 34 mm). 



SYSTEMATICS OF THE PLUSIINAE 



201 






57 




58 





60 



Figs 55-60 55, Ctenoplusia (Ctenoplusia) euchroa (ws = 41 mm). 56, 'Plusia (s.l.) megaloba (ws = 28 
mm). 57, Agrapha ahenea (ws = 37 mm). Note the well developed dorso-lateral hair tufts. 58, 
Plusiopalpa dichora (ws = 43 mm). 59, Argyrogramma verruca (ws = 31 mm). 60, Stigmoplusia 
chalcoides (ws = 37 mm) . 



202 



IAN J. KITCHING 




61 





63 






Figs 61-66 61, Eosphoropteryx thyatiroides (ws = 38 mm). 62, Polychrysia moneta (ws = 39 mm). Note 
the erect tegulae and scattered black scales on the forewing. 63, Chrysanympha formosa (ws = 33 mm). 
64, Panchrysia deaurata (ws = 37 mm). 65, P. v-argenteum (ws = 35 mm). 66, P. dives (ws = 26 mm). 



SYSTEMATICS OF THE PLUSIINAE 



203 








Figs 67-71 67, Euchalcia (Euchalcia) variabilis (ws = 38 mm). 68, 'Plusia' hampsoni (ws = 34 mm). 69, 
Pseudochalcia inconspicua (ws = 35 mm). 70, Desertoplusia bella (ws = 28 mm). 71, Euchalcia (Adeva) 
albavitta (ws = 27 mm). 



204 



IAN J. KITCHING 





73 



8 



74 







Figs 72-77 72, Rachiplusia nu (ws = 33 mm). 73, Erythroplasia rutulifrons (ws = 29 mm). 74, Loboplusia 
vanderweelei (holotype) (ws = 36 mm). Note the enlarged anal fold on the hindwing, the large dorsal 
hair tuft on the hindleg and the modified scales on the outer third of the hindwing. 75, Allagrapha aerea 
(ws = 40 mm). 76, Macdunnoughia (Macdunnoughia) confusa (ws = 36 mm). 77, M. (Puriplusia) 
purissima (ws = 32 mm) . 



SYSTEMATICS OF THE PLUSIINAE 



205 







■p 



78 








Figs 78-83 78, Sclerogenia Jessica (ws = 30 mm). 79, Antoculeora ornatissima (ws = 37 mm). 80, 
Autoplusia egena (ws = 40 mm). 81, A olivacea (ws = 43 mm). 82, 'Syngrapha' gammoides a" (ws = 46 
mm). 83, 'S. ' gammoides $ (ws = 44 mm). Note the sexual dimorphism in 'S. ' gammoides; the male is 
grey and more contrastingly patterned than the brown female. 



206 



IAN J. KITCHING 








88 




Figs 84-89 84, Diachrysia chryson (ws = 40 mm). 85, D. chrysitis (ws = 36 mm). 86, Anagrapha falcifera 
(ws = 35 mm). 87, Autographa gamma (ws = 40 mm). 88, 'Autographa' bractea (ws = 36 mm). 89, 
Cornutiplusia circumflexa (ws = 38 mm). 



SYSTEMATIC? OF THE PLUSIINAE 



207 



^mJJ* 



«6? <^>s 





91 



I 






fe.. 






^*SB 






r^GBB* 








■KM * . - 




Br '' ■ 
i 


W \ ^^ 


jST 


93 




1 \ 







Figs 90-95 90, Plusiafestucae (ws = 37 mm). 91 , P. venusta (ws = 33 mm). 92, Lophoplusia pterylota (ws 
= 38 mm). Note the strongly bipectinate antennae. 93, Syngrapha (Palaeographa) interrogationis (ws = 
33 mm). 94, S. (Syngrapha) devergens (ws = 28 mm). 95, S. (Caloplusia) hochenwarthi (ws = 28 mm). 



. 



208 



IAN J. KITCHING 







100 




\ 




Figs 96-101 96, Syngrapha (Syngrapha) parilis (ws = 26 mm). 97, Magusa orbifera (ws = 40 mm). 98, 
Stictoptera cucullioides (ws = 35 mm). Note the bulbous eyes. 99, Lophoptera sp. near squammigera (ws 
= 31 mm). 100, Paectes cristatrix (ws = 30 mm). Note the strongly bipectinate antennae. 101, Anuga \ 
constricta (ws = 36 mm). Note the very weakly bipectinate antennae. 



SYSTEMATICS OF THE PLUSIINAE 



209 





103 





105 





107 



Figs 102-107 102, Chalcopasta territans (ws = 36 mm). 103, Oncocnemis confusa (ws = 28 mm). 104, 
Calophasia lunula (ws = 29 mm). 105, Cucullia umbratica (ws = 50 mm). 106, Brachionycha sphinx (ws 
= 41 mm). 107, Diloba caeruleocephala (ws = 37 mm). 



210 



IAN J. KITCHING 









Figs 108-113 Probosces. 108, apex of proboscis, Autographa bractea. Note that the strengthening bars 
remain semicircular to the apex and that there are few apical sensilla. 109, apex of proboscis, Anuga 
constricta. Note that the strengthening bars break up apically into small plates and that there is a large 
number of apical sensilla. 110, styloconic sensillum, Autographa bractea. Ill, styloconic sensillum, 
Cucullia umbratica. 112, styloconic sensilla, Oncocnemis confusa. Note the longitudinal ridges that are 
absent in Autographa and Cucullia. 113, styloconic sensilla, Stictoptera cucullioides . Note the three 
poorly developed ventral ridges. Scale as indicated. 



SYSTEMATICS OF THE PLUSIINAE 



211 









Figs 114-119 Probosces. 114, styloconic sensillum, Paectes cristatrix. 115, styloconic sensillum, Anuga 
constricta. 116, styloconic sensillum, Paectes cristatrix. Note on Figs 114-116, the three enlarged 'wings'. 
117, apex of proboscis, Cucullia umbratica. Note that the strengthening bars are broken up apically into 
small nodules. 118, apex of proboscis, Anuga constricta. 119, base of proboscis, Anuga constricta. Note 
the regular arrangement of the microtrichia. Scale as indicated. 



212 



IAN J. KITCHING 




Figs 120-130 120-126, female frenula, right hindwings: (120) Anuga; (121) Paectes; (122) Stictoptera; 
(123) Diloba; (124) Autographa gamma; (125) Anadevidia; (126) Thysanoplusia. 127 ', 128, right 
hindwing apices: (127) Lamprotes; (128) Allagrapha. 129, 130, right forewing apices: (129) Pseudeva; 
(130) Lophoplusia. 



SYSTEMATICS OF THE PLUSIINAE 



213 




Figs 131-139 131-134, right hindwing venation: (131) Paectes; (132) Autographa gamma; (133) Cucullia; 
(134) Lophoptera. 135, 136, right forewing areoles: (135) Lophoplusia; (136) Lophoptera. 137-139, 
right hindwing venation: (137) Syngrapha (Syngrapha); (138) Chalcopasta; (139) Calophasia, vein M 2 is 
completely lost in this genus and is represented by a fold depicted as a dashed line. 



214 



IAN J. KITCHING 



o 



o 



o 



o° 


o o 


o 


O 


o 


o 


O 


o 

o 


o 
o 



o0 



o 



o 



o 



o 



o 



o 



o 



o 



140 




<S> o 



o o 
o 



o 



141 




Figs 140-146 140-142, scale base patterns: (140) Omorphina; (141) Sclerogenia; (142) Paectes. 143, 
tornal angle of Diloba, illustrating the apical remnant of vein CuP. 144-146, right pilifers, frontal view: 
(144) Stictoptera; (145) 'Autographa' bractea; (146) Diloba. Scale line for Figs 140-142 = 0-1 mm and for 
Figs 144-146 = 0-2 mm. 



SYSTEMATICS OF THE PLUSIINAE 



215 







150 





Figs 147-152 Heads. 147, dorsal view, Mouralia. 148, lateral view, Mouralia. 149, dorsal view, Anuga. 
150, lateral view, Anuga. 151, latero-frontal view, Syngrapha (Syngrapha). 152, frontal view, Stictop- 
tera. 



216 



IAN J. KITCHING 




Figs 153-159 153-157, labial palps: (153) Oncocnemis; (154) Panchrysia; (155) Anagrapha; (156) 
Chrysanympha; (157) Plusiopalpa. 158, 159, vom Rath's organ tubes: (158) Stictoptera; (159) Abrostola. 
Scale lines for Figs 153 and 155 = 0-5 mm and for Figs 158-159 = 01 mm. 



SYSTEMATICS OF THE PLUSIINAE 



217 




Figs 160-167 160-163, antennal segments: (160) Lophoplusia; (161) Paectes; (162) Eosphoropteryx; 
(163) Brachionycha. 164-167, left tegulae, dorsal view: (164) Abrostola; (165) Diloba; (166) Diachrysia; 
(167) Brachionycha. The broad dorsal arm rests latero-dorsally on the mesothorax while the apically 
microspinose ventral arm curves underneath the wings. 



218 



IAN J. KITCHING 




173 







h 



H 



Figs 168-177 168-170, ventral tegular arms, showing microspination: (168) Plusia; (169) Stictoptera; 
(170) Cucullia. 171-173, outlines of ventral tegular arms: (171) Syngrapha (Palaeographa); (172) 
Omorphina; (173) Paectes. 174, 175, outlines of dorsal tegular arms illustrating the marginal flange: 
(174) Agrapha; (175) Cucullia. 176, 177, patagia: (176) Anuga; (177) 'Autographa' bractea. The 
attachment point (dashed outline) is ventral and medial, with the narrower angle (uppermost in these 
drawings) directed laterally. Scale line for Figs 168-170 = 0-2 mm. 



SYSTEMATICS OF THE PLUSIINAE 



219 




178 







182 




183 




184 



Figs 178-184 178-181, male hind-basitarsi illustrating the basitarsal combs: (178) Plusiopalpa; (179) 
Thysanoplusia; (180) Ctenoplusia (Ctenoplusia); (181) Argyrogramma. 182-184, male fore-basitarsi: 
(182) Plusiotricha (Plusiotricha); (183) Stigmoplusia; (184) Argyrogramma. 



220 



IAN J. KITCHING 







188 





190 



Figs 185-190 185-187, hind-basitarsi: (185) Anagrapha; (186) Paectes; (187) Stictoptera. 188-190, 
hind-tibiae: (188) outer view, Syngrapha (Syngrapha); (189) outer view, Rachiplusia; (190) inner view, 
Mouralia. 



SYSTEMATICS OF THE PLUSIINAE 



221 





H 




196 




201 







199 



200 




Figs 191-204 191-194, fore-tarsi, third segment: (191) Lophoplusia; (192) Polychrysia; (193) Diachry- 
sia; (194) Anagrapha. 195, 196, fore-tibiae: (195) Rachiplusia; (196) Syngrapha (Caloplusia) 197-200 
outlines of hind-tibiae, tibial spurs and basitarsi: (197) Anuga; (198) Stictoptera; (199) 'Autographa' 
bractea; (200) Euchalcia (Euchalcia). 201-204, tarsal claws: (201) Cucullia; (202) Autographa gamma- 
(203) Plusiopalpa; (204) Oncocnemis. Scale line for Figs 201-204 = 0-5 mm 



222 



IAN J. KITCHING 




205 






207 






210 




214 





215 




208 



H 



Figs 205-215 205-207, clawed fore-tibiae: (205) Brachionycha; (206) Chalcopasta; (207) Oncocnemis. 
208, male St7, anterior edge uppermost, Anadevidia. 209, 210, lateral views from left of prothoracic 
furcae: (209) Lophoplusia; (210) Anuga. 211-213, prothoracic furcae and epimera, posterior views: 
(211) Lophoplusia; (212) Paectes; (213) Diloba. 214, 215, male St7s, anterior edges uppermost: (214) 
Stigmoplusia; (215) Argyrogramma. 



SYSTEMATICS OF THE PLUSIINAE 



223 





227 



228 



229 



Figs 216-229 216-221, internal flanges on the anterior edges of T2: (216) Cucullia; (217) Stictoptera; 
(218) Lophoptera; (219) Omorphina; (220) Abrostola; (221) Anagrapha. 222, 223, St2's, anterior edges 
uppermost: (222) Syngrapha (Caloplusia); (223) Paectes psaliphora Hampson. 224, 225, female T7's, 
posterior edges uppermost: (224) Adeva; (225) Autoplusia egena. 226-229, diagrammatic cross-sections 
of dorso-lateral scent patches on male T7's; scale arbitrary: (226) Agrapha; (227) Dactyloplusia; (228) 
Ctenoplusia (Ctenoplusia); (229) Argyrogramma. 



224 



IAN J. KITCHING 




230 




231 




Figs 230-234 Counter tympanal hoods and pleural pouches. The apodemes from St2 and the post- 
spiracular bars are shaded. 230, anterior view, RHS, Thysanoplusia orichalcea. 231, lateral view, RHS, 
Thysanoplusia orichalcea. 232, anterior view, LHS, Panchrysia. 233, anterior view, RHS, Stictoptera. 
234, anterior view, RHS, Calophasia. 



SYSTEMATICS OF THE PLUSIINAE 



225 





h 



H 



239 




Figs 235-240 St8 and sternal hair pencils, ventral views. 235, Brachionycha, hair pencils absent. 236, 
Cucullia, hair pencil represented by a sparse tuft of scales arising from a sclerotized line. 237, Magusa, 
the lateral arms of St8 have been flattened so that the hair pencils have been turned outwards through 
90°. 238, Paectes. 239, Stictoptera. 240, Anuga, the hair pencil pockets have been turned through 180° so 
as to face anteriorly instead of posteriorly. In Figs 237 and 238, and in subsequent illustrations of hair 
pencils, the pencils themselves are not drawn in detail but their limits are shown by dashed lines. If limits 
are not shown, this indicates that the hair pencil scales had become detached before drawing. In 
addition, the shading at the base of the pockets indicates the extent of the main fields of hair pencil scale 
insertions. 



226 



IAN J. KITCHING 




Figs 241-247 St8 and sternal hair pencils, ventral views. 241, Lamprotes. 242, Chrysodeixis (Chry- 
sodeixis) argentifera. Note the expanded secondary hair brushes, the spatulate apical scales and the 
complete fusion of the hair pencils and their pockets. 243, Macdunnoughia (Macdunnoughia) . 244, 
Autoplusia olivacea. 245, 'Autographa' bractea. 246, Plusiotricha (Eutheiaplusia). Note the reduced hair 
pencils and reduced secondary hair brushes. 247, Chrysodeixis (Pseudoplusia). Note the total loss of all 
hair pencils and brushes. 



SYSTEMAT1CS OF THE PLUSIINAE 



227 



248 




250 







h 



Figs 248-253 248, 249, St8 and hair pencil of Mouralia: (248) ventral view; (249) lateral view. 250-253, 
A8 and hair pencil structures of Chrysodeixis (Chrysodeixis) chalcites: (250) St8, posterior edge 
uppermost; (251) lateral view of A8 and hair pencil complex; note how the sagittate St8 supports the 
almost spherical hair brush ventrally, while T8 supports it dorsally, with the lateral arms of T8 curving 
laterally and ventrally to support the mouth of the hair brush pocket; (252) T8, posterior edge 
uppermost ; (253) anterior view of A8 and the hair pencil complex , snowing how the anterior part of St8 
supports the bases of the hair brushes. 



228 



IAN J. KITCHING 

h 




260 -L 



Figs 254-260 St8 and sternal hair pencils. 254, Trichoplusia, ventral view. 255, Trichoplusia, lateral view. 
256, Thysanoplusia, ventral view. 257, Thysanoplusia, lateral view. 258, Argyrogramma, ventral view; 
note the differentiation of the hair pencil scale insertions and the large semicircular flange of scales 
attached to the apex of the dorsal arms of St8. 259, Argyrogramma, anterior view of left hair pencil 
pocket, showing the different fields of hair pencil scale insertions, dorsal side is uppermost. 260, 
Argyrogramma, lateral view; note the long, thin, curved scales arising from the spatulate area of scale 
bases and projecting beyond the main hair pencils. 



SYSTEMATICS OF THE PLUSIINAE 



229 






264 



Figs 261-266 St8 and sternal hair pencils. 261 , Plusiotricha (Plusiotricha) , ventral view. 262 Plusiotricha 
(Plusiotricha), lateral view; note the large secondary hair brushes. 263, Ctenoplusia (Acanthoplusia) 
ventral view. 264, Ctenoplusia (Acanthoplusia), lateral view. 265, Plusiopalpa, ventral view 266' 
Plusiopalpa, lateral view. 



230 



IAN J. KITCHING 




Figs 267-271 St8 and sternal hair pencils. 267, Ctenoplusia (Ctenoplusia) , ventral view. 268, Ctenoplusia 
(Ctenoplusia), lateral view; note that the outer hair pencil scales are shorter than the inner scales. 269, 
Stigmoplusia, ventral view; note the large semicircular flange of scales attached to the dorsal arms of St8. 
270, Stigmoplusia, anterior view of left hair pencil pocket; note the differentiated fields of hair pencil 
scale bases. 271, Stigmoplusia, lateral view. 



SYSTEMATICS OF THE PLUSIINAE 



231 




Figs 272-278 272, 273, St8 and sternal hair pencils, Agrapha: (272) ventral view; (273) lateral view; note 
the differentiated hair pencils, similar to the condition seen in Argyrogramma (Figs 258-260). 274-279, 
unci: (274) Abrostola, lateral view; (275) Abrostola, dorsal view; (276) Anuga, lateral view; (277) 
Anuga, lateral view; (278) Mouralia, dorsal view; (279) Mouralia, lateral view. In Figs 274 and 279, the 
LAST'S are shaded, while the shading on Fig. 278 indicates the field of long, deciduous hairs that arise on 
the uncus of this genus. 



232 



IAN J. KITCHING 




Figs 280-286 Unci and tegumena. 280, Stigmoplusia, dorsal view. Note the lateral flanges that are unique 
to certain members of this genus. 281, Stigmoplusia, lateral view. 282, Oncocnemis, lateral view. 283, 
Syngrapha (Caloplusia) , lateral view . 284, Trichoplusia, lateral view. 285, Euchalcia (Euchalcia), lateral 
view. 286, Anuga, dorso-lateral view. Scale line for Fig. 286 = 0-5 mm. 



SYSTEMATICS OF THE PLUSIINAE 



233 




Figs 287-291 287-290, unci and tegumena: (287) Macdunnoughia (Macdunnoughia), lateral view; (288) 
Anadevidia, dorsal view; (289) Chrysodeixis (Chrysodeixis), lateral view; the LAST has been shaded; 
(290) Anagrapha, dorsal view. 291, lateral view of the terminal segments of Paectes, to illustrate the 
setose socii and the beak-like gnathos. The valves have been removed. 



234 



IAN J. KITCHING 




Figs 292-298 292, lateral view of the terminal segments of Anuga, to show the reduced uncus and large, 
curved, apically hooked socii. 293, 294, lateral views of unci and tegumena: (293) Argyrogramma; (294) 
Euchalcia (Adeva). 295-297, dorsal views of tegumena: (295) Erythroplasia; (296) Allagrapha; (297) 
Antoculeora. 



SYSTEMATICS OF THE PLUSIINAE 



235 




Figs 298-305 Vincula. 298, Agrapha, ventral view. 299, Stigmoplusia, dorsolateral view. 300, Stigmo- 
plusia, ventral view. 301, Thysanoplusia, ventral view. 302, Rachiplusia , ventral view. 303, Autographa 
gamma, ventral view. 304, Plusiotricha (Plusiotricha), ventral view. 305, Plusiotricha (Plusiotricha) , 
dorso-lateral view. 



236 



IAN J. KITCHING 




Figs 306-312 306, 307, pleurite of Ctenoplusia (Ctenoplusia), to illustrate its attachment to the vinculum 
and tegumen: (306) ventro-lateral view; (307) lateral view. 308-310, ventral view of vincula: (308) 
Chrysodeixis (Chrysodeixis); (309) Lophoptera; (310) Paectes. 311, 312, lateral views of vincula and 
pleurites: (311) Rachiplusia; (312) Anadevidia. 



SYSTEMATICS OF THE PLUSIINAE 



237 






Figs 313-321 Lateral views of pleurites and their attachments to their vincula and tegumena. 313, 
Mouralia. 314, Magusa. 315, Calophasia. 316, Antoculeora. 317, Oncocnemis. 318, Abrostola. 319^ 
'Autographa' bractea. 320, Diloba. 321, Argyrogramma. 



238 



IAN J. KITCHING 




Figs 322-334 322 , 323 , diagrammatic lateral views showing the relationship between the apex of the juxta 
and that of the aedeagus: (322) Abrostola; (323) generalized member of the Autoplusiina. 324-334, 
juxtae: (324) Mouralia, ventral view; (325) Abrostola, ventral view; (326) Sclerogenia, ventral view; 
(327) Sclerogenia, lateral view; (328) Dactyloplusia, ventral view; (329) Thysanoplusia, ventral view; 
(330) Plusiopalpa, ventral view; (331) Panchrysia, ventral view; (332) Chrysodeixis (Chrysodeixis) 
argentifera, lateral view; (333) Paectes, ventral view; (334) Chrysodeixis (Chrysodeixis) chalcites, ventral 
view. On Figs 326-331 and, subsequently, Figs 335-340, black shading at the base of the juxta indicates 
the area where it is fused to the valve flanges. 



SYSTEMATICS OF THE PLUSIINAE 



239 




Figs 335-346 335-341, juxtae: (335) Syngrapha (Palaeographa), ventral view; (336) Syngrapha 
(Palaeographa), lateral view; (337) Syngrapha (Syngrapha), ventral view; (338) Syngrapha (Caloplu- 
sia), ventral view; (339) Syngrapha (Caloplusia), lateral view; (340) Anagrapha, ventral view; (341) 
Chrysodeixis (Pseudoplusia), ventral view. 342, ventral view of the juxta and the bases of the valves of 
Agrapha, to illustrate the U-shaped juxta and the basal valve flanges. 343, base of the valves and juxta of 
Panchrysia, to show the bulge that appears between the valve bases when the valves are opened flat. 
344-346, postero-lateral views of transtillae: (344) Plusiopalpa; (345) Erythroplasia; (346) Euchalcia 
(Euchalcia) . 



240 



IAN J. KITCHING 




Figs 347-350 Left valves. 347, Abrostola. 348, Mouralia; note the apical twist to the valve, bearing a large 
fan of spatulate scales, the limits of which are indicated by the dashed line. 349, Cucullia. 350, 
Polychrysia. 



SYSTEMATICS OF THE PLUSIINAE 



241 




Figs 351-359 351-355, left valves: (351) Plusiotricha (Plusiotricha); (352) Plusiotricha (Eutheiaplusia); 
(353) Stigmoplusia; (354) Agrapha; the dashed line shows the limits of the short, curved, upright scales 
on the flattened edge of the sacculus; (355) Ctenoplusia (Acanthoplusia); the dotted oval indicates the 
position of the pad of inflated scales on the external surface of the valve. The small inverted 'u's on the 
apex of the valves in Figs 351-355 (and also on Figs 367 and 379) are the insertions of the basally directed 
apical scales found in the members of the Argyrogrammatini. The dark shading at the bases of several 
valves indicates to points of fusion with the juxta (cf. Figs 326-331 and 335-340). 356-359, clavi: (356) 
Oncocnemis; (357) Cucullia; (358) Antoculeora; (359) Sclerogenia. Scale line for Fig. 354 = 0-5 mm. 



242 



IAN J. KITCHING 




Figs 360-367 360-361, left valves: (360) Rachiplusia; (361) Sclerogenia. 362-365, subapical cornuti: 
(362) Euchalcia (Euchalcia), dorsal view; (363) Euchalcia (Euchalcia) , lateral view; (364) Euchalcia 
(Adeva), dorsal view; (365) Euchalcia (Adeva), lateral view. 366, left valve and juxta, Diachrysia. 367, 
left valve, Dactyloplusia. Scale lines for Figs 362-365 = 0-5 mm. 



SYSTEMATICS OF THE PLUSIINAE 



243 




Figs 368-371 Left valves. 368, Allagrapha. 369, Macdunnoughia (Puriplusia). 370, Macdunnoughia 
(Macdunnoughia); the black points are heavily sclerotized. 371, Loboplusia. 



244 



IAN J. KITCHING 




Figs 372-374 Left valves. 372, Autoplusia egena. 373, Autoplusia olivacea. 374, 'Syngrapha' gammoides. 
Note in the last species that the juxta is fused with the clavus, which has lost its connection with the valve 
completely. 



SYSTEMATICS OF THE PLUSIINAE 



245 




Figs 375-378 375-377, left valves: (375) Paectes; (376) inner view of Antoculeora; (377) dorsal view of 
Antoculeora. 378, both valves and juxta of Erythroplasia; note the asymmetrical harpes. 



246 



IAN J. KITCHING 




Figs 379-385 379-382, left valves: (379) Trichoplusia; (380) Syngrapha (Caloplusia); (381) Autographa 
gamma; (382) Lophoplusia. 383-385, auricular cornuti; (383) Loboplusia; (384) Macdunnoughia 
(Macdunnoughia); (385) Eosphoropteryx. Scale lines for Figs 380 and 383-385 = 0-5 mm. 



SYSTEMATICS OF THE PLUSIINAE 



247 




Figs 386-396 386, 387, left valves: (386) Syngrapha (Palaeographa); (387) Stictoptera. 388, 389, vincula 
and valves, outer view: (388) Paectes; (389) Euchalcia (Euchalcia). 390-392, diagrammatic lateral views 
of aedeagi illustrating the three states of character 157: (390) state 0; (391) state 1 ; (392) state 2. 393-395, 
diagrammatic ventral views of aedeagi to illustrate the three states of character 158: (393) state 0; (394) 
state 1 ; (395) state 2. Scale for Figs 390-395 is arbitrary. 396, dorsal view of the basal part of the vesica of 
Pseudeva, to illustrate the patch of curved cornuti. 



248 



IAN J. KITCHING 




Figs 397-402 Aedeagi and vesicas. 397, Chrysodeixis (Pseudoplusia), lateral view. 398, Mouralia, 
ventral view. 399, Mouralia, lateral view. 400, Plusiopalpa adrasta, ventro-lateral view. 401 , Dactyloplu- 
sia, lateral view. 402, Syngrapha (Palaeographa). The dashed ovals on Figs 400 and 402 represent the 
opening of the cuticular simplex. The dashed lines on Figs 397 and 401 represent the internally 
invaginated sacs. 



SYSTEMATICS OF THE PLUSIINAE 



249 




409 



410 







f- 



H 



Figs 403-413 403-405, subapical cornuti: (403) 'Autographa bractea; (404) Plusia; (405) Lophoplusia. 
406, diagrammatic representation of the serpentine-coiled vesica of the genus Plusiotricha; scale 
arbitrary. 407, Abrostola, aedeagus, ventral view. 408, 409, subapical cornuti: (408) Chrysanympha; 
(409) Panchrysia. 410, 411, aedeagi: (410) Ctenoplusia (Acanthoplusia) , lateral view; (411) Plusidia, 
lateral view. 412, 413, anal papillae and intersegmental membrane: (412) Anadevidia, showing the 
membranous pouch anterior to the anal papillae; (413) Cucullia, showing the unexpanded membrane 
anterior to the anal papillae. 



250 



IAN J. KITCHING 




Figs 414-420 Anal papillae. 414, Ctenoplusia (Acanthoplusia) . 415, Thysanoplusia. 416, Anadevidia, 
417, Paectes, posterolateral view. 418, Lophoptera, postero-lateral view. 419, Panchrysia. 420, 
Cucullia. 



SYSTEMATICS OF THE PLUSIINAE 



251 






426 




Figs 421-430 421-426, female segments A8: (421) Abrostola, ventral view; (422) Abrosto la, lateral view; 
(423) Mouralia, lateral view; (424) Plusiopalpa, lateral view; (425) Anagrapha, lateral view; (426) 
Paectes, lateral view. 427, 428, antra: (427) Syngrapha ( Palaeographa) , ventral view; (428) Rachiplusia, 
ventral view. 429, 430, female segments A8, lateral views: (429) Anuga; (430) Euchalcia (Adeva). 



252 



IAN J. KITCHING 




Figs 431-437 Antra. 431, Macdunnoughia (Macdunnoughia) , ventral view. 432, Macdunnoughia (Puri- 
plusia), ventral view. 433, Sclerogenia, ventral view, illustrating the tuft of erect scales at the mouth of 
the antrum. 434, Plusidia, ventral view. 435, Lamprotes, ventral view. 436, Antoculeora, lateral view, 
right. 437, Antoculeora, ventral view; note the large, asymmetrical, sclerotized, open antrum. 



SYSTEMATICS OF THE PLUSIINAE 



253 




444 



Figs 438-444 438-440, A8 and antrum of Autoplusia olivacea: (438) postero-ventral view; (439) ventral 
view; (440) lateral view, left. 441, dorsal view of the flattened section with recurved edges of the ductus 
bursae of Macdunnoughia (Macdunnoughia). 442-444, ducti and corpora bursae: (442) Dactyloplusia, 
ventral view; (443) Argyrogramma, lateral view; (444) Agrapha. 



254 



IAN J. KITCHING 




Figs 445-448 Ducti and corpora bursae. 445, Rachiplusia. 446, Thysanoplusia. 447, Trichoplusia. 448, 
Plusiotricha (Eutheiaplusia) . 



SYSTEMATICS OF THE PLUSIINAE 



255 




Figs 449-453 Ducti and corpora bursae. 449, Allagrapha. 450, Macdunnoughia (Macdunnoughia). 451, 
Macdunnoughia (Puriplusia) . 452, Eosphoropteryx. 453, Erythroplasia. 



256 



IAN J. KITCHING 




Figs 454-456 Ducti and corpora bursae. 454, Autoplusia. 455, Sclerogenia. 456, Antoculeora. The dashed 
line indicates the discontinuity in ornamentation of the corpus bursae (character 212). 



SYSTEMATICS OF THE PLUSIINAE 



257 




Figs 457-460 Ducti and corpora bursae. 457, distal part of the corpus bursae of Paectes, illustrating the 
concave signum and the circular ridges. 458, Cornutiplusia, the dashed lines show the extent to which the 
ductus sclerotization continues onto the apex of the corpus bursae. 459, Anagraphs 460, Lophoptera. 



258 



IAN J. KITCHING 






Figs 461-465 461-463, ducti and corpora bursae: (461) Plusia; (462) Lophoplusia; (463) 'Autographa' 
bractea. 464, 465, corpora bursae: (464) Calophasia, showing the sclerotized longitudinal ridge; (465) 
Lophoptera, illustrating the invaginated, spinose signum. 



SYSTEMATICS OF THE PLUSIINAE 

Index 



259 



Abrostula 101 102 103 140 177 178 

180 190 
abrostolina 141 190 
Abrostolini 84 87 140 175 179 
abrostoloides 124 
abstrusu 166 
acalypta 128 134 149 
Acanthoplusia 1 13 133 147 171 
accenlifera 139 147 
acuta 92 132 143 145 172 190 
/t^va97 107 115 151 152 174 
adiaphora 147 
adrasla94 148 
aemula&\ 162 

aenofusa 128 129 134 135 149 
aerea79 91 155 
aereoides 85 155156160 
«g/u»«81 147 149 171 
/lgrap/i«79 113 146 
ahenea 79 146 
fli/i 166 167 
albavitta 152 

albostriata 81 128 134 144 148 177 
a/ias 166 

Altagrapha 79 106 117 155 
u«w« 128 134 149 
alpina 165 
aZ/era 164 

«///Vo/« 133 166 180 
amp/a 79 162 
amy dm 147 
Anadevidia 1 19 142 153 160 177 

178 190 
Anagrapha 103 106 1 15 159 161 

164 
angulidens 133 166 
angulum 92 
Anigraea 169 
anophioides 141 178 
Antoculeora 93 107 117 119 158 

177 
antsalova 149 
/*««£« 94 124 169 
arachnoides 92 128 
argentifera 93 95 132 143 179 
Argyrogramma 107 113 148 
Argyrogrammatini 84 87 142 175 

179 
argyrosigna 81 
asclepiadis 141 
augusta 136 150 
aurantiaca 140 
aurata 150 
aureus 150 
Autographa 1 15 156 157 161 162 

177 180 
,4«top/i«ifl93 117 154158 
Autoplusiina 153 175 

W/ M cfl85 155 160 
basigera 91 128 149 
M/a 172 
bellieri 152 



/)/£'/( 142 156 160 

biloba!9 9[ 161 

bimaculata 161 

bi partita 145 

fovea 125 164 

Brachionycha94 108 119 169 180 

foractoj81 115 162 177 

brassicae 145 

brevi.striata 144 

buraetica 180 

c-aureum 153 

calceolaris 146 

californica 79 161 180 

«////'.s7« 128 

Calophasia 94 105 119 

Caloplusia 107 1 15 154 164 166 

Caloplusiini 87 

camptogamma 1 48 

camptosema 1 56 

carcassoni 97 1 46 

cashmirensis 151 152 

cefea79 165 166 177 

Cerviphtsia 190 

c/ifl/ci/es 93 96 132 143 145 

chalcoides 149 

Chalcopasta 94 103 105 108 119 

cheiranthi 152 

chinghaiensis 152 

chlorocharis 152 

ChrysanymphaW 103 124 151 177 

Chrysaspidia 190 

chrysitis 7883 124 139 155 160 

( hrysodeixis 93 95 102 1 1 1 143 172 

179 190 
cAryson 83 106 160 
Chrysoptera 190 
circumflexa 92 162 
circumscripta HI 149 
compos ita 166 167 
con^tM 156 157 190 
consona 152 
contextu 163 
coreae 160 

Cornutiplusia 106 115 162 
corusca 79 1 62 
crassisigna 135 156 157 
cristalrix 94 
cry plica 166 
Ctenoplusia 113 133 145 147 148 

177 
Cwbaw 190 
Cucut Ha 94 100 101 106 107 112 

119 
cuprea 152 

Dactyloplusia 102 143 145 

da/e; 143 

<fowM81 124 144 145 

deaurata 78 151 

Desertoplusia 172 

devergens 125 133 165 166 167 180 

Diachrysia 83 102 155 159 178 



diasema 125 161 164 
dichora9? l 103 148 180 
D(7ofe« 94 103 109 119 169 
diversigna 125 164 
rf/V«125 151 
</o/era 128 
dorsiflava 136 151 
dyscapna 146 

egena93 129134139159 

egenella 159 

egenoides 158 

ekekei 145 

emmetra 156 

Eosphoropteryx 150 

epargyra 134 

epigaea 164 

epistilba 149 

eriosoma 143 

Erythroplasia 1 17 154 177 178 190 

esmeralda 150 151 

Euchalcia 106 1 15 151 152 174 177 

180 
Euchalciina 149 153 175 
Euchalciini 87 
eugrapha 147 
Eutelia 169 

Euteliinae 119 167 168 169 
Eutheiaplusia 97 101 102 108 146 
excelsa 162 
exequialis 190 
exquisila 125 128 

falciferaSS 161 
festucae 78 S3 139 163 
flagellum 161 

jlammifera 165 
fletcheri 97 146 

//«ri/i« 128 145 
formosa 78 151 
furahatai 190 
furcifera 147 

gamma 115 139 145 161 
paranoides 93 134 154 159 

gtrrfa 78 90 152 
giffardi 164 
gilarovi 165 
gorilla 144 
gromieri 128 

hainanensis 149 
hampsoni 151 
hebetata 134 142 
herbuloti 147 
hildebrandti 148 
hochenwarthi 125 133 166 167 
homoia 145 

ichinosei 147 
/gnea 125 133 165 166 167 
ignicollis 145 
tlluminata 93 132 



260 



IAN J. KITCHING 



illustrata 159 
imperatrix 124 152 
impulsa 143 

includens 91 139 143 145 
inconspicua 151 174 
indicator 124 
Inguridia 190 
intermixta 81 144 145 
interrogationis 165 180 
italica 136 152 

Jessica 158 
y'oto 162 

kaszabi 140 
tefofl 93 96 132 
kostjuki 162 

Lamproto 106 107 115 153 

latistigma 147 

lavendula 78 

lenzeni 151 

/eonwa 124 142 155 156 160 

leucostigma 128 134 

limbirena92 114 124 128 147 

livida 97 146 

Looop/ws/a 94 97 117 157 

/£>«</?/« 93 158 

longisigna 144 

Lophoplusia 103 115 163 

Lophoptera 94 101 103 124 168 169 

lushanensis 158 

Macdunnoughia 117 154 156 161 

177 190 
Magt«a94 119 167 
major 141 
mapongua 128 
mappa 79 162 
megaloba 135 
megista 134 149 
melanocephala 124 
Melanoplusia 190 
meretricia 146 
metallica 162 
micans 124 

microgamma 125 133 166 167 
mikadina 153 
Mimanuga 169 
minor 158 
minutus 144 
mishanensis 151 
moneta 150 151 
monosigna 156 157 
montana 125 133 165 166 167 
morigera 150 180 
Mouralia 107 141 177 178 180 

nade/a 106 155 160 
Neoplusia 190 
nz 91 96 140 144 145 
nichollae 163 
Nigramma 127 
nigh burs a 94 171 
nigrisigna 162 
«u 154 



nubeculosa 125 
nyiwonis 165 180 

obtusisigna 92 

ochreata 81 144 145 

octoscripta 164 

Odontodes 127 169 

olivacea 93 134 159 

Omorphina 97 103 140 178 

Omorphinini 87 140 175 

Oncocnemis 94 103 107 108 119 

orichalcea 83 92 128 139 145 

ornata 151 

ornatissima 81 83 93 158 

orophila 125 164 

ottolenguii 164 180 

oh 91 154 

oxy gramma 85 91 128 134 144 148 

Paecto 94 107 169 

paghmana 172 

Palaeographa 102 103 115 164 165 

180 
Palaeoplusia 190 
palligera 124 153 
Panchrysia 151 
paraplesia 149 
Paraplusia 190 

panfo 78 124 125 133 165 166 167 
pasiphaeia 161 
Penicillaria 169 
peponis 134 142 
Perloplusia 190 
p/iocea 128 134 
photeina 128 

P/wj/a 83 107 115 163 177 190 
'Plusia' 87 152 
Plusidia 115 152 
Plusiina 153 159 175 
Plusiinae 138 

Plusiini 85 87 142 149 175 179 
Plusiopalpa 93 101 103 108 113 148 

180 
Plusiotricha 97 107 1 13 133 146 147 
Podioplusia 190 
pokhara 94 157 
polvcampta 124 
Polychrysia 103 106 107 150 152 

178 180 
polydamia 190 
pram 146 
precationis 79 161 
pseclrocera 78 164 
Pseudeva 115 153 
Pseudochalcia 152 174 
pseudogamma 161 
Pseudop/usza 113 143 179 
pseudopyropia 190 
pterylota 78 164 177 
pulchrina 139 162 180 
Puriplusia 117 156 
purissima 156 
purpurigera 153 
putnami 163 
pyrenaica 165 
pyropia 81 124155 



Rachiplusia 107 108 117 121 153 

166 
rectangula 79 165 166 177 
rectilinea 128 
rubida 161 
rutulifrons 81 124 155 

sackenii 125 133 164 
sansoni 125 161 
schalisema 156 
Sclerogenia 107 117 157 177 
Scleroplusia 190 
selecta 164 
semirosea 145 
semivitta 156 
separanda 152 
serraticornis 90 152 
sestertia 92 128 
Shensiplusia 94 171 
sAisa93148 
5/ca 78 150 
siderifera 152 
sigillata 147 
szgna/a 81 128 148 149 
snowii 125 164 
sphinx 180 
splendida 152 
squammigera 94 
stenochrysis 160 
Sticloptera 94 103 112 168 169 
Stictopterinae 119 167 168 
Stigmophisia 97 107 108 113 149 
subaerea 149 
subchalybaea 128 
surena 165 

Syngrapha 115 161 162 164 166 
177 180 

tancrei 151 
tarassota 81 128 147 
Targalla 169 
letragona 156 
tetrastigma 92 
thaumasia 148 
thyatiroides 150 
Thysanoplusia 107 145 
tibetana 125 
tine to ides 141 
transbaikalensis 165 
Trichoplusia 102 107 144 
trigemina 140 141 
t hop us 180 
fr/p/os/a 139 140 141 
Zz/Z/z 160 

u-aureum 165 
t/nca 190 

v-alba 161 
v-argenteum 151 
vanderweelei 94 157 
variabilis 139 150 152 
venusta 163 190 
vermiculata 134 147 



SYSTEMATICS OF THE PLUSIINAE 261 

verruca 91 148 149 vittata 92 128 148 180 Yerongponga 190 

viettei 128 145 

violacea 164 177 wukongensis 190 

virgula 125 154 166 Zonoplusia 107 144 145 178 

viridisigma 164 xanthoides 152 zosimi 124 160 



V. 



British Museum (Natural History) 

Milkweed butterflies: their cladistics and biology 

P. R. Ackery & R. I. Vane-Wright 

The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of 
their biology have stimulated far more attention than can be justified by species numbers 
alone. In recent years, an expansive literature has grown, considering aspects of their 
courtship and pre-courtship behaviour, migration, larval hostplan associations, mimicry and 
genetics. The popularity of danaines among biologists can certainly be attributed to this 
combination, within one small group, of so many of the factors that make butterflies such an 
interesting group to study. The obvious need to place this wealth of biological data within an 
acceptable systematic framework provided the impetus for this volume. 

Started eight years ago within the conventions of evolution by natural selection and 
Hennig's phylogenetic systematics, the book is now largely about natural history (what the 
animals have and do, where they live and how they develop) and natural groups - as revealed 
by a form of analysis approaching that practised by the new school of 'transformed cladistics'. 
The authors have prepared a handbook that will appeal to a wide range of biologists, from 
museum taxonomists to field ecologists. 

425 pp, 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive bibliography. 
ISBN 565 00893 5. 1984. Price £50. 






Titles to be published in Volume 54 



Studies on the Old World species of Holothrips (Thysanoptera: Phlaeothripidae) 

By S. Okajima 

Spectacles and Silver Ys: a synthesis of the systematics, cladistics and biology of the Plusiinae 
(Lepidoptera: Noctuidae) 

By I. J. Kitching 

A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr 
(Hymenoptera: Formicidae) 

By B.Bolton 



Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk 
Printed in Great Britain by Henry Ling Ltd, Dorchester